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Thescelosaurus

Thescelosaurus is a genus of small to medium-sized herbivorous neornithischian dinosaurs belonging to the family Thescelosauridae, known from the Maastrichtian stage of the Late Cretaceous period (approximately 72–66 million years ago) in western North America.^[1] These bipedal ornithopods, characterized by their conservative morphology and elongated skulls, measured up to 4.1 meters in length and weighed around 340 kilograms in adulthood, inhabiting coastal plain environments such as swamps and river channels.^[2] Fossils indicate a diet primarily consisting of low-lying vegetation, supplemented possibly by roots and tubers detected through acute olfaction, with evidence suggesting some semi-fossorial behaviors like burrowing.^[2]^[1] The genus was established by Charles W. Gilmore in 1913 with the type species T. neglectus, based on specimens from the Lance Formation in Wyoming, including a partial skeleton that highlighted its robust build and dental adaptations for grinding plant matter.^[1] Two additional valid species are recognized: T. garbanii from the Hell Creek Formation in Montana, distinguished by differences in tarsal anatomy, and T. assiniboiensis from the Frenchman Formation in Saskatchewan, the smallest of the trio at around 3 meters long.^[3]^[1] Other proposed species, such as T. edmontonensis, have been synonymized or reclassified, reflecting ongoing taxonomic refinements based on comparative anatomy.^[3] Recent studies, including CT scans of cranial material, have revealed specialized neuroanatomy in T. neglectus, such as enlarged olfactory bulbs comprising about 3% of the brain volume and an elongated anterior semicircular canal for enhanced balance, supporting inferences of ecological adaptations like foraging in low-light or buried substrates just prior to the Cretaceous–Paleogene extinction event.^[2] With numerous specimens documented from formations like the Hell Creek and Scollard, Thescelosaurus provides key insights into the diversity and survival strategies of small-bodied dinosaurs in the final ecosystems of the Mesozoic era.^[3]^[1]

Discovery and history

Initial discovery of T. neglectus

The initial fossils attributed to Thescelosaurus neglectus were collected prior to its formal description, beginning with a paratype specimen (USNM 7758) unearthed in 1889 by Orlando A. Peterson from the Lance Formation at Lance Creek, Converse County, Wyoming.^[4] This material included several vertebrae, scapulae, and elements of the fore and hind limbs.^[4] The type specimen (USNM 7757), a more complete partial skeleton comprising articulated vertebrae, ribs, a partial pelvis, and most limb bones but lacking the skull and neck vertebrae, was discovered in July 1891 by John Bell Hatcher and William H. Utterback from the nearby type locality at Doegie Creek, also in Converse County, Wyoming, within the same Late Cretaceous Lance Formation.^[4] These specimens remained unstudied for over two decades until paleontologist Charles W. Gilmore examined them at the United States National Museum, recognizing their significance as a new taxon.^[4] In a preliminary publication in Smithsonian Miscellaneous Collections, Gilmore formally named the genus and species Thescelosaurus neglectus on May 24, 1913, designating USNM 7757 as the holotype.^[4] The generic name Thescelosaurus derives from the Greek words theskelos (marvelous or wonderful) and sauros (lizard), while the specific epithet neglectus (Latin for neglected or overlooked) reflects the long delay in describing the material despite its collection over 20 years earlier.^[4] Gilmore's initial description portrayed T. neglectus as a small, bipedal ornithopod dinosaur approximately 3.5–4 meters in length, provisionally placing it within the family Camptosauridae based on postcranial resemblances to larger ornithopods like Camptosaurus.^[4] In a more detailed osteological study published in 1915 in the Proceedings of the United States National Museum, Gilmore revised this classification, aligning T. neglectus with the Hypsilophodontidae due to shared traits such as single-headed posterior thoracic ribs and the phalangeal formula of the manus, emphasizing its status as a primitive ornithopod with agile, bird-like hindlimbs adapted for terrestrial locomotion. These early works established T. neglectus as a representative of the diverse small-bodied ornithischians from the latest Cretaceous of western North America.

Subsequent species and revisions

In 1940, Charles M. Sternberg named Thescelosaurus edmontonensis based on a partial skeleton (holotype CMN 8537) consisting of vertebrae, pelvis, hind limbs, and other elements, collected from the Horseshoe Canyon Formation (then part of the Edmonton Formation) in Alberta, Canada.^[5] This material was distinguished from the type species T. neglectus primarily by its more gracile build and differences in pelvic structure, including a narrower prepubis.^[5] The genus expanded further with the description of Thescelosaurus garbanii by William J. Morris in 1976, based on a fragmentary postcranial skeleton (holotype LACM 33542) including cervical and dorsal vertebrae, partial pelvis, femur, and foot elements from the Hell Creek Formation in Montana.^[6] Morris highlighted key anatomical differences, such as preserved vertebral elements (five posterior cervical and 11 anterior dorsal vertebrae) and more robust limb proportions, with a relatively shorter tibia relative to the femur, suggesting adaptations for a larger body size approaching 4 meters in length.^[6] Peter M. Galton conducted a major revision of Thescelosaurus in 1974, examining multiple specimens and proposing classifications within ornithopod dinosaurs, which laid groundwork for later taxonomic adjustments.^[7] By the early 1980s, Galton further refined these views, synonymizing T. edmontonensis with Parksosaurus warreni due to overlapping vertebral and limb features, while questioning the validity of T. garbanii owing to its fragmentary nature and potential overlap with T. neglectus in robusticity.^[7] These revisions emphasized differences in vertebral counts and hind limb ratios as diagnostic, but noted challenges in distinguishing species from ontogenetic or individual variation. Mid-20th century debates centered on whether specimens attributed to Thescelosaurus represented a single conservative genus or multiple distinct ones, particularly after Sternberg erected Parksosaurus in 1937 for material previously considered T. warreni, based on differences in forelimb proportions and overall slenderness.^[5] These discussions influenced early understandings of ornithopod diversity in the Late Cretaceous, with revisions suggesting a more unified taxonomy under fewer genera.^[7] Such taxonomic shifts had brief phylogenetic implications, positioning Thescelosaurus as a basal neornithischian with conservative traits bridging earlier hypsilophodontids and more derived forms.

Bugenasaura and exceptional specimens

In 1995, paleontologist Peter M. Galton erected the genus Bugenasaura for two specimens recovered from the Hell Creek Formation (late Maastrichtian) of South Dakota, recognizing them as distinct from Thescelosaurus neglectus due to their more robust cranial features. The holotype (SDSM 7210) consists of a partial skull, including left and right dentaries with teeth, maxillae, and other fragments, while the paratype (SDSM 40488) is an isolated right dentary; both exhibit prominent ridges on the maxilla and dentary interpreted as sites for muscular cheek attachments.^[8] The type species B. infernalis was also designated to include the holotype of Thescelosaurus garbanii (LACM 33542), a fragmentary postcranial skeleton originally described in 1976, based on similarities in jaw morphology and dental structure.^[9] Subsequent analysis in 2009 by Clint A. Boyd and colleagues led to the synonymization of Bugenasaura with Thescelosaurus neglectus, arguing that the diagnostic traits proposed by Galton—such as the robust dentary and specific tooth morphology—fall within the range of intraspecific variation observed in T. neglectus specimens from the same formation. Specifically, shared features include the asymmetrical enamel on teeth (thicker lingually), the form of the predentary articulation, and the overall jaw architecture, which do not warrant generic separation. This revision emphasized the conservative morphology of basal neornithischians and reduced the number of valid genera in the Hell Creek ornithopod assemblage.^[10] One of the most notable Thescelosaurus specimens is NCSM 15728, a well-preserved juvenile skeleton discovered in 1993 from the Hell Creek Formation in Harding County, South Dakota, by paleontologist William R. Hammer; measuring about 3 meters in length, it includes a nearly complete skull, articulated vertebrae, ribs, and partial limbs, providing key insights into ontogeny. In 2000, Paul E. Fisher and coauthors reported a reddish, grapefruit-sized mass in the thoracic cavity as a fossilized four-chambered heart, based on CT imaging that revealed internal structures resembling ventricles and major vessels, suggesting advanced cardiovascular anatomy. This interpretation was challenged in 2001 by Timothy Rowe and colleagues, who applied higher-resolution CT scans and concluded the structure was not organic tissue but an inorganic ironstone concretion formed by sediment diagenesis, with its shape and position resulting from postmortem deformation and mineralization processes rather than anatomical preservation. Their analysis highlighted the rarity of soft-tissue fossilization and the need for rigorous verification, as similar concretions occur in other Hell Creek matrix. The "Willo" specimen (named after the discoverer's wife's nickname) remains significant for its skeletal completeness, despite the debunked heart claim.

Recent finds including T. assiniboiensis

In 2011, paleontologists Caleb M. Brown, Clint A. Boyd, and Dale A. Russell described a new species of Thescelosaurus, T. assiniboiensis, based on a nearly complete skeleton collected from the late Maastrichtian Frenchman Formation in Saskatchewan, Canada. The holotype specimen, RSM P 1225.1, represents an adult individual and includes partial cranial material along with most of the postcranial skeleton, providing the first substantial skull elements for the genus beyond fragmentary remains.^[11] This species is distinguished from the earlier T. neglectus and T. garbanii by its larger overall size—approximately 20% longer than the T. neglectus holotype—and unique features in the pelvic girdle, such as a more elongate preacetabular process of the ilium and a straighter ischium shaft that retains plesiomorphic traits similar to those in the related taxon Parksosaurus. Cranially, T. assiniboiensis exhibits autapomorphic conditions absent in prior species, including a supraoccipital bone with a prominent midline foramen and a broader quadrate with a distinct ventral notch, enhancing understanding of morphological variation within the genus.^[11] Since the early 2000s, tentative reports of teeth from the Campanian Judith River Formation in Montana were initially assigned to Thescelosaurus but later reclassified to related taxa such as Orodromeus.^[12] These finds indicate potential broader distribution for the genus in pre-Maastrichtian deposits but require further assessment.^[13] A 2023 computed tomography (CT) study by David J. Button and Lindsay E. Zanno examined the skull of T. neglectus (specimen NCSM 15728, an adult from the Lance Formation) to reconstruct its endocranial anatomy, revealing specialized sensory capabilities such as an enlarged olfactory region comprising about 3% of the endocast volume for enhanced smell and an elongated anterior semicircular canal for superior balance detection, though its hearing range was restricted to low frequencies (296–2150 Hz).^[2] This analysis did not propose any new species but underscored ecological adaptations in basal neornithischians without altering taxonomic boundaries.^[2] In 2024, a new thescelosaurine, Fona herzogae, was described from the Cenomanian Cedar Mountain Formation in Utah, suggesting burrowing behaviors in early members of the family and providing insights into the evolution leading to Thescelosaurus.^[14] Despite these advances, knowledge of Thescelosaurus remains incomplete, particularly regarding skull material, as only fragmentary cranial elements were confidently referred to T. neglectus prior to recent studies, limiting detailed comparisons across species. Recent 2020s phylogenetic analyses of Thescelosauridae suggest potential undiscovered Asian relatives, given comparisons between North American taxa like Thescelosaurus and East Asian forms such as Koreanosaurus boseongensis, indicating possible faunal exchanges across Beringia in the Late Cretaceous.^[2]

Description

Cranial anatomy

The skull of Thescelosaurus is characteristically low and elongated, exhibiting typical ornithopod features such as a toothless beak at the rostral tip and a diastema between the predentary and maxilla.^[1] In adults, the overall skull length measures approximately 30–40 cm, based on composite measurements from well-preserved specimens like NCSM 15728, where the premaxilla reaches 101.4 mm, the maxillary tooth row 90.4 mm, and the dentary 146.7 mm.^[1] The cranium is triangular in dorsal view, with a slight transverse crushing in some examples, and portions of nearly every bone are represented in the most complete known skull (NCSM 15728).^[1] Dentition is a key diagnostic trait, featuring leaf-shaped teeth with thick, asymmetrical enamel and marginal denticles suited for grinding vegetation.^[1] The premaxilla bears 5–6 teeth per side, while the maxilla and dentary each hold up to 20 tooth positions (e.g., 20 in NCSM 15728 and 18 in TLAM.BA.2014.027.0001), with crowns displaying fine ridges and a constriction at the base.^[1] A prominent horizontal ridge runs along the maxilla, accompanied by coarse, obliquely inclined ridges, representing an autapomorphy of the genus.^[3] The braincase is relatively derived, resembling that of Dryosaurus and Dysalotosaurus, with features like a groove on the prootic and a shared fossa for cranial nerves VII and VIII.^[1] Orbits are large and rugose-margined, bordered by the prefrontal, lacrimal, jugal, and postorbital, indicating enhanced visual capabilities.^[1] Narial openings are positioned rostrally, enclosed by the premaxilla and nasal bones.^[1] Species variations include differences in T. assiniboiensis, which possesses broader premaxillae compared to T. neglectus, along with a convex posterior squamosal margin and a unique median foramen on the supraoccipital.^[11] Complete skulls are rare, with most cranial material fragmentary; for instance, the former genus Bugenasaura (now synonymous with Thescelosaurus) is based primarily on dentaries exhibiting pronounced ridges interpreted as cheek muscle attachments.^[3]

Postcranial skeleton

The postcranial skeleton of Thescelosaurus exhibits a robust body plan indicative of basal neornithischians, with adults reaching lengths of 3–4.1 meters and estimated masses up to 340 kg.^[3]^[2] The animal was bipedal, supported by powerful hindlimbs, but retained robust forelimbs compatible with facultative quadrupedality.^[15] The axial skeleton comprises approximately 9 cervical vertebrae, 16 dorsal vertebrae, 5 sacral vertebrae, and around 50 caudal vertebrae.^[16] Neural spines are low and thin across the presacral series, forming rectangular plates that increase slightly in height posteriorly.^[16] The caudal series features haemal arches (chevrons) beginning at the second vertebra, providing structural support to the stiffened tail reinforced by ossified tendons.^[16] Forelimbs are robust, with the humerus measuring approximately 60% of femur length (e.g., humerus 215 mm, femur 355 mm in the holotype).^[16] Hindlimbs display a straight femur and a slender, straight fibula closely appressed to the tibia posterodistally, facilitating bipedal progression.^[16] The pelvis includes an ilium with a pronounced, elongate preacetabular process that tapers to a triangular cross-section and exceeds the postacetabular portion in length.^[16] Ribs are double-headed anteriorly and single-headed posteriorly, with a broad thoracic cage; ossified tendons, interpreted as intercostal plates, occur laterally along the anterior dorsal ribs, with up to 7 preserved per side in some specimens.^[16]^[17] These plates may have contributed to postural stability.^[17] Among species, T. garbanii differs in possessing relatively longer metacarpals compared to T. neglectus.^[3]

Skin and integument

Skin impressions from Thescelosaurus specimens in the Lance Formation reveal small, polygonal scales on the tail and flanks, resembling the non-overlapping scales of modern lizards. These impressions indicate a reptilian-style integument, with scales averaging a few millimeters in diameter and arranged in an irregular pattern without larger osteoderms or tubercles dominating the surface. Impressions from the holotype (USNM 7751) also show similar small, non-overlapping scales on the tail.^[16] No feathers or filamentous structures have been identified in any Thescelosaurus fossils, consistent with the scaly texture preserved. Comparisons to the closely related Parksosaurus, another late Maastrichtian neornithischian, support an overall non-feathered integument, with similar scale morphologies inferred from shared postcranial features and environmental context.29[0325:TROTBN]2.0.CO;2/Taxonomic-Revision-of-the-Basal-Neornithischian-Taxa-Thescelosaurus-and/10.1671/0272-4634(2009)29[0325:TROTBN]2.0.CO;2.full) The limited fossil evidence suggests possible sparse osteoderms along the dorsal surface, though this interpretation is tentative and based on comparisons with other basal ornithopods; such structures, if present, would have provided minimal armor rather than extensive plating. Skin preservation is exceptionally rare in Thescelosaurus, occurring in fewer than 5% of known specimens due to taphonomic biases favoring bone over soft tissue in fluvial and floodplain deposits.^[16]

Classification and systematics

Valid species and synonyms

The genus Thescelosaurus encompasses three valid species according to current taxonomic consensus: the type species T. neglectus, T. garbanii, and T. assiniboiensis.^[14] The type species T. neglectus was established by Charles W. Gilmore in 1913, based on the holotype (USNM 7757, a partial skeleton lacking the skull) and paratype (USNM 7758, preserving additional postcranial elements including previously unrecognized cranial material) from the late Maastrichtian Lance Formation in Niobrara County, Wyoming.^[18]^[3] It is diagnosed by features such as participation of the calcaneum in the midtarsal joint and five cranial autapomorphies, including frontals wider at the midorbital level.^[3] T. garbanii was named by William J. Morris in 1976 from the holotype (LACM 33542), a fragmentary postcranial skeleton including vertebrae, ribs, and hindlimb elements from the Hell Creek Formation in Garfield County, Montana.^[3] Its validity was confirmed in a 2009 revision through shared apomorphies with other Thescelosaurus species but distinguished by the exclusion of the calcaneum from the midtarsal joint and other postcranial differences, supported by phylogenetic analysis.^[3] The youngest species, T. assiniboiensis, was described by Caleb M. Brown and colleagues in 2011 based on the holotype (RSM P 1225.1), an articulated partial skeleton preserving much of the skull, axial skeleton, pelvic girdle, and hindlimbs from the late Maastrichtian Frenchman Formation in Saskatchewan, Canada.^[11] It is approximately 13% smaller than the T. neglectus holotype and diagnosed by two cranial autapomorphies: a convex dorsal and posterior margin of the squamosal, and a distinct median foramen on the supraoccipital connecting the myelencephalon to the dorsal surface; the preserved pelvic girdle further supports distinction through plesiomorphic traits like the ilium shape, more similar to outgroups than to other Thescelosaurus species.^[11] Several proposed taxa have been relegated to synonymy or invalidity. The genus Bugenasaura, erected by Peter M. Galton in 1995 (with a detailed description in 1997) for the holotype (SDSM 7210), a partial dentary from the late Maastrichtian Hell Creek Formation in South Dakota, was synonymized with Thescelosaurus in 2009 due to indistinguishable dentary morphology from T. neglectus and insufficient unique diagnostic features, rendering Bugenasaura infernalis a nomen dubium.^[19]^[3] Similarly, T. edmontonensis, named by Charles M. Sternberg in 1940 from isolated postcranial elements in the late Maastrichtian Scollard Formation of Alberta, Canada, is regarded as a junior subjective synonym of T. neglectus owing to overlapping diagnostic traits without unique autapomorphies.^[3] Species validity within Thescelosaurus relies on demonstrable autapomorphies confirmed through comparative anatomy and phylogenetic analysis, alongside adequate skeletal material to rule out ontogenetic or intraspecific variation; for example, T. assiniboiensis was upheld despite its smaller size because its autapomorphies persist across growth stages and differ consistently from T. neglectus and T. garbanii.^[3]^[11] The 2009 revision emphasized that taxa like Bugenasaura and T. edmontonensis lack such distinguishing characters when directly compared to the type species.^[3] Debates persist on subtle biogeographic differentiation, with T. assiniboiensis from Canadian deposits exhibiting some plesiomorphic features (e.g., in the pelvic girdle) that may reflect population-level variation between northern and southern North American assemblages, potentially warranting further scrutiny of geographic isolation in late Maastrichtian thescelosaurines.^[11] Overall, Thescelosaurus is represented by multiple specimens—over a dozen well-preserved partial to near-complete skeletons—predominantly of T. neglectus from U.S. formations, underscoring its relative abundance in terminal Cretaceous ecosystems.^[2]

Phylogenetic relationships

Thescelosaurus is recognized as a basal euornithopod within the clade Thescelosauridae, where cladistic analyses consistently position it as the sister taxon to Parksosaurus, with Orodromeus as the next closest relative in a polytomy or sequential branching among North American Cretaceous neornithischians.^[10] This placement emphasizes the conservative morphology of thescelosaurids as early-diverging ornithopods, distinct from more derived groups like iguanodontians.^[20] Updated phylogenetic matrices from the 2020s, such as those in Madzia et al. (2021), recover Thescelosaurus as a non-hadrosauromorph ornithopod, with its lineage diverging from the stem leading to advanced ornithopods around 100 million years ago during the Early Cretaceous.^[20] Supporting this position are key synapomorphies of Thescelosauridae, including a pronounced notch on the posterior margin of the premaxilla and a thin, medially projecting flange on the fibula that articulates with the tibia.^[10] The 2024 description of Fona herzogae, a basal thescelosaurine from the Cenomanian Mussentuchit Member of the Cedar Mountain Formation in Utah, reinforces this framework by recovering it as the earliest known member of Thescelosaurinae and a potential ancestor to Thescelosaurus species, highlighting the early evolution of burrowing adaptations in the lineage as evidenced by associated postcranial modifications shared with the semi-fossorial Oryctodromeus.^[14] Alternative hypotheses from some 2010s phylogenetic studies, however, propose a closer affinity to Dryosauridae, nesting Thescelosaurus nearer to Jurassic taxa like Dryosaurus within a broader basal ornithopod radiation rather than as a distinct thescelosaurid lineage.^[21]

Paleobiology

Locomotion and posture

Thescelosaurus was primarily a facultative biped, capable of both bipedal and quadrupedal stances, as indicated by its skeletal proportions and forelimb morphology. The forelimbs were relatively short, with a humerus length of approximately 215 mm compared to a femur length of 355 mm, yielding a humerus-to-femur ratio of about 0.6, which allowed the forelimbs to contact the ground during a quadrupedal stance for activities such as foraging while maintaining the ability for bipedal progression.^[16] This configuration is consistent with basal ornithopods, where the forelimbs could be positioned with elbows tucked in and palms facing medially, enabling ground contact but rendering quadrupedal locomotion awkward and unlikely habitual due to limited humeral range of motion and medial palm orientation that did not facilitate forward propulsion.^[22]^[23] Estimated speeds for Thescelosaurus, based on limb proportions and comparisons to small bipedal ornithopods, suggest it could achieve running velocities of 20–40 km/h, though larger individuals were likely restricted to walking or slow trotting gaits around 15–20 km/h.^[24] While direct trackways attributable to Thescelosaurus are unknown, stride analyses from related thescelosaurine or basal ornithopod ichnites support moderate speeds in this range for similar-sized animals, reflecting adaptations for evasion rather than sustained high-speed pursuit.^[24] The robust hind limbs, with a femur longer than the tibia, further indicate a build suited for stability over speed, prioritizing endurance in varied terrains.^[16] The tail of Thescelosaurus functioned as a counterbalance during bipedal locomotion, stiffened by extensive ossified tendons extending from the dorsal, sacral, and caudal regions to maintain rigidity and prevent lateral sway.^[16] This stiffening, comprising rod-like structures along the tail's length (which accounted for about half the total body length), aided in upright bipedal posture with a horizontal neck orientation for stability during movement.^[16] Biomechanical assessments of basal ornithopod forelimb and girdle morphology highlight how such proportions provided stability on uneven terrain, allowing Thescelosaurus to navigate forested or riverine environments without compromising balance.^[25]

Sensory abilities and behavior

A computed tomography (CT) scan of the skull of Thescelosaurus neglectus specimen AMNH 5060 reveals specialized sensory capabilities, particularly in olfaction. The olfactory bulbs comprise approximately 3% of the total endocast volume, a relatively large proportion exceeding that of most ornithischians and overlapping with values seen in extant rodents and lagomorphs, indicating an acute sense of smell adapted for locating buried food resources such as roots and tubers or detecting predators from a distance.^[2] Hearing in Thescelosaurus appears to have been limited, with the inner ear structures supporting a narrow frequency range of roughly 296–2150 Hz and a best sensitivity around 1100–1200 Hz, comparable to crocodilians but reduced relative to other dinosaurs like Dysalotosaurus, potentially impairing detection of high-pitched predator vocalizations or environmental cues.^[2] The tympanic region, featuring a rod-shaped stapes extending to the paroccipital process, aligns with this moderate auditory capacity.^[26] Large orbits preserved in cranial specimens further suggest that vision played a key role, likely enabling diurnal activity and visual foraging in forested environments.^[26] The reptile encephalization quotient (REQ) of Thescelosaurus falls at or below the average for reptiles of similar body mass, with cerebral hemispheres occupying about 30% of the endocast—less than in hadrosaurids (~40%)—implying relatively modest cognitive abilities suited to simpler social structures and smaller group sizes compared to other Late Cretaceous ornithischians.^[2] Behavioral inferences point to a semi-fossorial lifestyle, with neurological features like the enlarged olfactory bulbs and elongated anterior semicircular canals providing enhanced balance for navigating underground tunnels, complemented by robust forelimbs and strong claws for excavation and refuge construction.^[2] This burrowing adaptation, representing the first such specialization identified in ornithischian dinosaurs, likely served as protection from predators and extreme surface conditions, and is corroborated by the 2024 description of Fona herzogae, an early Cenomanian thescelosaurine ancestor exhibiting pronounced semi-fossorial traits such as reinforced shoulder girdles and enlarged olfactory regions.^[14]

Intercostal plates and other features

Intercostal plates in Thescelosaurus consist of thin, dorsoventrally expanded, D-shaped bony structures positioned laterally to the anterior dorsal ribs, with the first plate typically associated with the second rib and subsequent plates overlapping adjacent ribs and one another.^[27] These plates form through endochondral ossification and are interpreted as ossified intermuscular septa that provided attachment sites for hypaxial musculature, as evidenced by the presence of Sharpey's fibers.^[27] The primary function of the intercostal plates likely involved biomechanical stabilization of the ribcage and torso during locomotion, potentially enhancing respiratory efficiency by facilitating rib movements analogous to uncinate processes in birds, though not homologous to them.^[27] This arrangement may have supported an active lifestyle by aiding lung ventilation through costal mechanics.^[27] These plates are present across all valid species of Thescelosaurus, though their number varies ontogenetically and by specimen preservation; for instance, up to seven plates have been documented in a mature individual of T. neglectus (NCSM 15728).^[27] They are more commonly preserved in articulated skeletons of mature animals, suggesting ossification occurred later in development.^[27] Other notable features include the dental morphology, with maxillary and dentary teeth featuring 15–20 longitudinal ridges that converge in a crescentic pattern toward the crown base, accompanied by lingual wear facets indicative of propalinal jaw action for processing tough vegetation.^[28] This wear pattern supports a herbivorous diet focused on abrasive plant matter.^[28] Intercostal plates represent an evolutionary novelty within Neornithischia, absent in basal ornithischians such as Haya griva, and exhibit convergence with rib-associated structures like avian uncinates in derived theropods, likely evolving independently to enhance thoracic support.^[27]^[29]

Controversial soft tissue preservation

In 1993, a nearly complete skeleton of Thescelosaurus neglectus (specimen NCSM 15728, nicknamed "Willo") was discovered in the Hell Creek Formation of South Dakota, and subsequent analysis revealed an iron-rich structure within the thoracic cavity. In 2000, researchers described this structure as a fossilized four-chambered heart, resembling those of mammals or birds rather than reptiles, based on medical imaging that suggested vascular features and a divided aorta. This interpretation implied the presence of an advanced cardiovascular system in ornithopod dinosaurs, potentially indicating higher metabolic rates and endothermy in non-avian dinosaurs. However, subsequent examinations refuted the heart claim. A 2001 computed tomography (CT) scan by Rowe and colleagues revealed the structure to be a mineral concretion composed of iron-cemented sandstone, lacking diagnostic cardiac anatomy such as atria, ventricles, or coronary arteries, and showing features consistent with inorganic formation rather than soft tissue preservation. Similar concretions have been documented in other Hell Creek Formation fossils, where they form pseudomorphs around voids or decayed tissues, mimicking biological structures due to diagenetic processes. A 2011 reexamination, including histological analysis of a fragment from the concretion, confirmed it consisted of cemented sand grains with no organic biomarkers or soft tissue remnants, further supporting its identification as an abiotic artifact. Soft tissue preservation in Thescelosaurus remains rare, with no verified examples of internal organs beyond the debated "Willo" case. A 2023 CT study of Willo's skull reconstructed lost soft tissues like the brain and inner ear but confirmed the absence of preserved organs elsewhere in the specimen, underscoring the challenges of exceptional preservation in ornithischian dinosaurs.^[2] The scientific consensus views the "heart" as a geological concretion rather than biological tissue, highlighting taphonomic biases in the Hell Creek Formation where rapid burial and mineral-rich sediments favor such pseudofossils over true soft tissue mineralization.

Paleoecology

Geological and geographical distribution

Thescelosaurus is known exclusively from the Late Cretaceous of North America, with all confirmed fossils dating to the Maastrichtian stage approximately 72 to 66 million years ago.^[1] Although thescelosaurine ornithopods, the clade to which Thescelosaurus belongs, have earlier records—such as the Cenomanian species Fona herzogae from about 99 million years ago in Utah—no definitive Thescelosaurus specimens predate the Maastrichtian.^[14] Fossils of Thescelosaurus have been recovered from several formations across western North America, spanning the United States and Canada. In the western United States, specimens occur in the Hell Creek Formation of Montana, North Dakota, and South Dakota; the Lance Formation of Wyoming; and the Laramie Formation of Colorado within the Denver Basin.^[1] In Canada, remains are documented from the Scollard Formation of Alberta and the Frenchman Formation of Saskatchewan.^[1] The stratigraphic range of the genus is confined to the Maastrichtian stage, spanning formations from the Hell Creek and Lance to the Frenchman Formation, reflecting a distribution tied to the latest Cretaceous continental deposits of Laramidia.^[11] No confirmed Thescelosaurus fossils have been reported from Asia or Europe, despite the potential for faunal exchange across the Bering land bridge during the Late Cretaceous, which facilitated dispersal of other dinosaur groups between North America and Asia.^[30]

Taphonomy and abundance

Fossils of Thescelosaurus are predominantly preserved as disarticulated skeletons within fluvial mudstone deposits of the Hell Creek and Lance Formations, reflecting transport and scattering in riverine environments prior to burial.^[31] Rare articulated specimens, such as the nearly complete T. assiniboiensis holotype from the Frenchman Formation, indicate episodes of rapid burial that protected remains from significant post-mortem disturbance.^[2] Exceptional preservation includes occasional soft tissue impressions, as seen in the "Willo" specimen (NCSM 15728), where iron concretions formed around decaying organic material, mimicking internal structures but later confirmed as mineralization artifacts rather than original tissues.^[32] In Maastrichtian faunas, Thescelosaurus represents a common element, comprising approximately 8% of identified dinosaur specimens in unbiased surveys of the Hell Creek Formation and 5-10% of microvertebrate assemblages recovered through screen-washing techniques at sites like those in Montana.^[31] This relative abundance positions it as one of the more prevalent small ornithischians, occasionally outnumbering hadrosaur remains in localized microfaunal samples from floodplain deposits.^[32] Preservation biases affect the fossil record of Thescelosaurus, with juvenile individuals underrepresented due to the fragility of small bones, which are more susceptible to erosion and destruction during weathering and transport in fluvial settings.^[31] Bonebeds containing multiple juvenile Thescelosaurus alongside other taxa, such as those from the "convenience store" locality in the Hell Creek Formation, suggest mass mortality events, possibly linked to flooding or drought, followed by concentrated deposition.^[2] Over 40 specimens of Thescelosaurus have been collected, primarily during the 20th century from key sites in the Lance and Hell Creek Formations, with the genus first described in 1913 based on material gathered in 1891.^[26] Modern paleontological surveys, employing systematic screen washing, continue to recover mostly isolated bone fragments and teeth, enhancing understanding of intraspecific variation but yielding fewer complete skeletons.^[31] Thescelosaurus fossils occur up to the Cretaceous-Paleogene (K-Pg) boundary, including a disarticulated leg from the Tanis site in North Dakota preserved within impact-related sediments, marking it among the last known non-avian dinosaurs in North America.^[2] Features suggestive of burrowing behavior in some specimens raise the possibility of localized refugia that may have buffered against immediate extinction pressures, though no post-boundary survival is documented.^[33]

Paleoenvironments and fauna

Thescelosaurus inhabited subtropical floodplains and coastal plains characterized by meandering rivers and seasonal streams draining into the Western Interior Seaway during the late Maastrichtian stage of the Late Cretaceous.^[34] These environments featured lush, multi-tiered vegetation dominated by angiosperms, thermophilous conifers, ferns, and broad-leaved deciduous trees, supporting a diverse herbivorous community.^[35] The dinosaur preferred habitats near river channels, as evidenced by its prevalence in sandstone deposits indicative of channel margins and levees, where it likely foraged on low-lying vegetation.^[34] Paleoclimatic reconstructions for these regions indicate a warm-temperate to subtropical climate with mean annual temperatures of approximately 20–25°C, marked by wet conditions punctuated by dry seasons.^[36] Evidence from paleosols and foliar physiognomy supports moderate seasonality without freezing winters, fostering the growth of thermophilous flora adapted to humid, riverine settings.^[37] As a low-browser herbivore, Thescelosaurus occupied a niche involving the consumption of ferns, horsetails, and understory angiosperms, using its dental battery for shearing tough plant material; it competed for resources with larger hadrosaurs like Edmontosaurus but partitioned space by favoring channel-proximal areas over distal floodplains.^[34] Predation pressure came primarily from tyrannosaurids such as Tyrannosaurus rex, prompting behavioral adaptations including possible semi-fossorial habits for refuge.^[2] In broader faunal assemblages from formations like the Hell Creek and Lance, Thescelosaurus coexisted with ceratopsians (e.g., Triceratops), hadrosaurs (e.g., Edmontosaurus), ankylosaurians, pachycephalosaurians, and small theropods, forming part of a rich, multi-tiered ecosystem on the eve of the Cretaceous–Paleogene extinction.^[34] Neurological specializations in its braincase suggest enhanced sensory capabilities suited to burrowing, potentially aiding predator avoidance and exploitation of burrow microhabitats amid this diverse fauna.^[2] As a generalist herbivore with versatile ecological tolerances, Thescelosaurus persisted as one of the last non-avian dinosaurs until the asteroid impact approximately 66 million years ago, highlighting its adaptability in a fluctuating end-Cretaceous landscape.^[35]

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