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Ceratops

Ceratops is a dubious genus of herbivorous ceratopsid dinosaur that lived during the Late Cretaceous period (Campanian stage), approximately 76 million years ago, in what is now western North America.^[1]^[2] The genus was established by paleontologist Othniel Charles Marsh in 1888 as the type genus of the family Ceratopsidae, with the single species Ceratops montanus, based on fragmentary holotype fossils (USNM 2411) consisting of a pair of postorbital horncores and a partial occipital condyle recovered from the Judith River Formation in Fergus County, Montana.^[1]^[2] The name Ceratops derives from the Greek words keras (horn) and ops (face), reflecting the prominent horned cranial structure typical of ceratopsids.^[1] As a member of Ceratopsidae, Ceratops would have been a large, quadrupedal ornithischian dinosaur characterized by a robust beak-like mouth for cropping vegetation, a bony frill extending from the back of the skull, and epiossified (bony) projections on the frill, though no frill material is preserved for this genus.^[2] The known fossils indicate straight, elongate postorbital horncores that are mediolaterally compressed and lack distinctive features, measuring about 20 cm in length and showing a roughened surface texture consistent with mature individuals.^[2] Due to the limited and non-unique morphology of these remains—which closely resemble horncores of other centrosaurine ceratopsids such as Albertaceratops and Kosmoceratops—Ceratops montanus is classified as a nomen dubium, meaning its generic distinctiveness cannot be confidently confirmed.^[2] Subsequent research has occasionally referred additional isolated elements, such as teeth from the Judith River Formation and the equivalent Oldman Formation in Alberta, Canada, to Ceratops, but these assignments are tentative and do not resolve the genus's taxonomic ambiguity.^[2] The Judith River Formation, a fluvial and floodplain deposit spanning about 79 to 75 million years ago, has yielded a diverse ceratopsid assemblage including well-known genera like Centrosaurus, Chasmosaurus, and Judiceratops, highlighting a period of rapid evolutionary radiation among horned dinosaurs in Laramidia (the western landmass of North America).^[2] Despite its uncertain status, Ceratops represents an early contribution to the understanding of ceratopsian diversity and the "Bone Wars" rivalry between Marsh and Edward Drinker Cope, which accelerated dinosaur discoveries in the late 19th century.^[1]

Discovery and history

Initial discovery

The genus Ceratops was first established in 1888 based on fossils collected by John Bell Hatcher during a field expedition for Othniel Charles Marsh in the Judith River Formation of Fergus County, Montana, USA.^[3] The holotype specimen (USNM 2411), housed at the Smithsonian Institution's National Museum of Natural History, comprises a partial occipital condyle and a pair of supraorbital horn cores from Late Campanian sediments approximately 77–75 million years old.^[4] Marsh promptly described and named the type species Ceratops montanus in December of that year, recognizing it as representative of a primitive horned dinosaur and establishing the family Ceratopsidae, from which the clade Ceratopsia derives its name. This hasty naming occurred amid the Bone Wars, the fierce rivalry between Marsh and Edward Drinker Cope that spurred rapid publications of incomplete material to claim scientific priority over emerging dinosaur discoveries.^[5]

Species and nomenclature

The genus Ceratops was established by Othniel Charles Marsh in 1888 with the naming of the type species C. montanus, based on the holotype specimen USNM 2411—a fragmentary partial skull consisting of two supraorbital horn cores (each approximately 22 cm long) and an occipital condyle—collected from the Campanian-age Judith River Formation of Fergus County, Montana. The material was discovered earlier that year by field collector John Bell Hatcher and represents the first named ceratopsid genus, though its extreme fragmentation limited diagnostic utility from the outset. In 1889, Marsh proposed a second species, Ceratops horridus, based on a more complete partial skull (YPM 1821) from the Lance Formation of Wyoming, but he promptly reassigned it as the type species of the new genus Triceratops upon recognizing additional distinguishing features, such as a third nasal horn. Marsh also named C. alticornis that year for isolated horn cores (YPM 1820) from the Denver Formation of Colorado, later synonymized with Triceratops horridus. These early designations reflected the rapid taxonomic flux during the Bone Wars era, where Marsh and rival Edward Drinker Cope competed to classify emerging ceratopsian fossils; Cope had described related fragmentary material under Monoclonius (e.g., M. canadensis in 1888 and M. recurvicornis in 1889), some of which was later considered for referral to Ceratops. Subsequent nomenclatural efforts included John Bell Hatcher's 1905 revision, which transferred three Monoclonius species to Ceratops—C. canadensis, C. recurvicornis, and C. plicocompsus—based on perceived similarities in horn morphology, though these were provisional and lacked new material. Earlier tentative links to Cope's Agathaumas (1872, based on caudal vertebrae from the Pierre Shale) and Polyonax mortuarius (1874, based on a partial skeleton including limb bones from Colorado) were proposed but rejected due to insufficient overlap with Ceratops type material and their own fragmentary conditions. Under International Code of Zoological Nomenclature (ICZN) rules, Ceratops is now classified as a nomen dubium because the holotype of C. montanus provides no autapomorphic features sufficient to distinguish it from other ceratopsids, rendering the genus and type species indeterminate. This assessment was formalized in modern reviews, including Penkalski and Dodson (1999), who emphasized the meager, non-diagnostic nature of the remains and noted potential affinities to primitive chasmosaurines like Avaceratops without confirming referral.^[6] No complete or referred specimens reliably support the genus today, confining it to historical taxonomic context.

Description

Skull morphology

The holotype of Ceratops (USNM 2411) consists of a pair of postorbital horncores and a partial occipital condyle, providing limited insight into its cranial morphology. The horncores are approximately 20-22 cm long, subtriangular at the base, and curve outward sharply, with a roughened surface texture indicating mature individuals.^[3] Due to the fragmentary nature of the remains and the genus's status as a nomen dubium, detailed skull features cannot be confidently attributed to Ceratops. Tentatively referred material, such as isolated squamosals (e.g., USNM 4802), suggests a short, broad frill comparable to other Judith River ceratopsids, but these assignments remain uncertain.^[3] No frill, preorbital region, rostrum, or dental elements are preserved in the holotype, precluding descriptions of primitive traits like fenestrae or beak structure.

Postcranial skeleton

No postcranial remains are known for Ceratops montanus. Due to the limited holotype material and taxonomic ambiguity, body size and posture cannot be determined with confidence. Comparisons to contemporary ceratopsids suggest it was a medium-sized quadruped, but specific estimates are unavailable.

Classification

Taxonomy

Ceratops is classified within the Linnaean hierarchy as follows: Kingdom Animalia, Phylum Chordata, Class Reptilia, Order Ornithischia, Suborder Cerapoda, Infraorder Ceratopsia, Family Ceratopsidae, Genus Ceratops.^[2] The genus serves as the eponymous basal taxon and type genus for the infraorder Ceratopsia, which was formally established by Othniel Charles Marsh in 1890 based on fragmentary material initially collected by Edward Drinker Cope. Marsh named the type species Ceratops montanus in 1888 from postorbital horncores and an occipital condyle recovered from the Judith River Formation in Montana. Due to the limited and non-diagnostic nature of the holotype material, Ceratops montanus is regarded as a nomen dubium, with the postorbital horncores potentially attributable to other ceratopsids such as Spiclypeus shipporum, Albertaceratops nesmoi, or Kosmoceratops richardsoni.^[2] Despite this, the genus Ceratops is retained in taxonomic nomenclature for its historical significance as the foundational taxon of Ceratopsia and its utility in cladistic analyses of basal ceratopsians.^[7] Marsh briefly described a second species, Ceratops paucidens, in 1890, transferring material originally assigned to the hadrosaur Hadrosaurus paucidens (1889). However, subsequent studies have reaffirmed that this material belongs to a hadrosaurid dinosaur and is not ceratopsian.^[8] By the early 20th century, workers such as Barnum Brown refined the understanding of Ceratops montanus as a primitive member of Ceratopsia distinct from more derived ceratopsids.^[7]

Phylogenetic relationships

Due to the extremely fragmentary nature of the holotype material—limited to a pair of postorbital horncores and a partial occipital condyle—no detailed phylogenetic analysis has included Ceratops montanus. The straight, elongate horncores resemble those of several derived ceratopsids from the Late Cretaceous of Laramidia, including Albertaceratops (Centrosaurinae), Kosmoceratops, and Spiclypeus (Chasmosaurinae), suggesting an uncertain but likely position within Ceratopsidae (Neoceratopsia).^[2] Its Late Campanian age (~76 Ma) precludes a basal position within Ceratopsia, as more primitive ceratopsians are known from the Jurassic and Early Cretaceous of Asia. Ceratops shares a North American distribution with other Late Cretaceous ceratopsids but lacks close relatives among earlier Asian forms, consistent with the radiation of Neoceratopsia in Laramidia following migration from Asia. Its occurrence in the Judith River Formation highlights the early diversification of ceratopsids, though its exact relationships remain unresolved due to the non-diagnostic fossils.^[2]

Paleobiology

Diet and feeding

Given its nomen dubium status and fragmentary remains, the paleobiology of Ceratops is poorly known and must be inferred from other centrosaurine ceratopsids. As a ceratopsid, Ceratops was an herbivorous dinosaur that engaged in low-level browsing, consuming vegetation such as ferns, cycads, horsetails, and angiosperms available in its Late Cretaceous (Campanian) floodplain environment.^[9] This reflects the adaptations of ceratopsids to process tougher, fibrous plant material using a complex dental battery, rather than the softer foliage exploited by basal ceratopsians.^[9] The feeding apparatus of ceratopsids included a multi-layered dental battery with leaf-shaped teeth arranged in functional rows along the jaw, enabling precise orthal occlusion and transverse grinding through palinal motion for efficient processing of vegetation.^[9] This system, combined with a robust beak-like rostrum for cropping, supported a powerful masticatory apparatus suited to the available flora.^[10] Biomechanical analyses of ceratopsids indicate substantial bite forces, amplified by elongated tooth rows, tall coronoid processes, and strong adductor muscles, allowing processing of tough, fibrous plants—far exceeding the capabilities of basal forms.^[10] Studies suggest ceratopsids were restricted to low browsing heights of around 1 meter, despite their large body size (up to 6-9 meters in length), due to neck posture and skull orientation.^[9] As a centrosaurine ceratopsid, Ceratops likely exploited a range of low vegetation, with its inferred horns and frill potentially aiding in defense during feeding. This niche contributed to ceratopsid diversification within herbivorous communities in Laramidia.^[2]

Growth and ontogeny

Due to the limited fossils, growth and ontogeny in Ceratops are inferred from related ceratopsids, such as centrosaurines like Pachyrhinosaurus. Bone histology reveals rapid early growth driven by fibrolamellar bone tissue, enabling juveniles to reach subadult sizes quickly.^[11] Growth rates in ceratopsids supported attainment of adult body lengths of 5-7 meters by late adolescence. Evidence for sexual dimorphism in ceratopsids is limited, with some studies suggesting subtle cranial differences, but no confirmed patterns in horn or frill development from available fossils.^[12] Lifespan estimates for ceratopsids range from 20-30 years, with maturity reached around 10-15 years based on lines of arrested growth (LAGs) and growth curve analyses in taxa like Triceratops.^[13] Later stages show slower growth with parallel-fibered bone and an external fundamental system (EFS), indicating determinate growth.^[11] Ontogenetic changes in ceratopsids include closure of cranial fenestrae and elaboration of horns and frills with age, as seen in Protoceratops and Triceratops series, reflecting maturation of defensive and display structures.^[12]

Paleoecology

Habitat and distribution

Ceratops is known solely from the Judith River Formation in north-central Montana, United States, where its holotype and limited additional material were recovered. This formation consists of nonmarine deposits laid down in a fluvial-deltaic setting, featuring meandering rivers, extensive floodplains, and associated wetlands such as swamps and oxbow lakes.^[14]^[15] The temporal range of Ceratops spans approximately 77 to 75 million years ago during the Late Campanian epoch, aligning with the Judithian North American Land Mammal Age; all known specimens derive from this interval within the formation, with no records from outside western North America.^[16]^[17] The paleoenvironment of the Judith River Formation supported a warm, humid climate conducive to lush vegetation, including dominant conifers such as Sequoia and Thuja, ferns like Osmunda, and understory elements including horsetails, alongside emerging angiosperms such as Populus and Quercus.^[14]^[18] This setting, part of a coastal plain in the Western Interior foreland basin, featured moderate to high precipitation that fostered herbivore-supporting ecosystems.^[19] Fossils of Ceratops, including cranial elements, are primarily preserved in cross-bedded channel sandstones and associated overbank deposits, indicating post-mortem transport via fluvial currents followed by rapid burial to minimize disarticulation and weathering.^[20]^[21]

Contemporaries and interactions

Ceratops inhabited the Judith River Formation of Montana during the Campanian stage of the Late Cretaceous, approximately 77–75 million years ago, where it coexisted with a diverse assemblage of dinosaurs. Among herbivores, it shared its environment with hadrosaurids such as Gryposaurus and Kritosaurus, which dominated the fauna as large browsers and grazers, as well as ankylosaurids including Euoplocephalus (formerly Palaeoscincus) and Edmontonia, known for their armored bodies and low-slung postures. Predatory theropods were represented by tyrannosaurids like Daspletosaurus, which preyed on medium- to large-sized dinosaurs, and smaller carnivores such as Troodon, a troodontid that likely scavenged or hunted juveniles.^[22]^[23]^[16] As an early ceratopsid, Ceratops occupied a mid-sized herbivorous niche, estimated at around 5 meters in length, functioning primarily as a browser of low-lying vegetation in floodplain and riverine habitats. This role likely overlapped with that of ornithopods like Gryposaurus, potentially leading to competition for understory plants such as ferns and cycads in the forested undergrowth of Laramidia's coastal plains. Unlike more derived ceratopsids, there is no fossil evidence, such as bonebeds indicative of mass mortality in herds, suggesting Ceratops lived solitarily or in small family groups rather than large social herds.^[16]^[24] Predation pressures on Ceratops were significant, given its primitive morphology lacking prominent horns or a robust frill for defense, making it vulnerable to attacks from apex predators like Daspletosaurus. This vulnerability is inferred from bite marks and pathologies observed on bones of contemporary and related ceratopsians, including tooth-scored squamosals and healed injuries consistent with tyrannosaurid encounters, which often targeted the vulnerable flanks or necks of herbivores. Such interactions highlight Ceratops' position lower in the food web compared to heavily armored or gregarious later ceratopsids.^[23]^[16] Within the broader biodiversity of Laramidia, the Western Interior landmass of Late Cretaceous North America, Ceratops represented a relict primitive form persisting amid a rapid radiation of more advanced neoceratopsians. The Judith River Formation preserves at least six valid ceratopsid species, including centrosaurines like Avaceratops, Albertaceratops, and Lokiceratops, and chasmosaurines such as Judiceratops, Medusaceratops, and the transitional Spiclypeus, illustrating a high diversity of horned dinosaurs evolving complex cranial ornamentation for display and defense. This assemblage underscores Ceratops' role as an early, unspecialized member of a clade that diversified dramatically in the Campanian, filling varied ecological roles in a dynamic, river-dominated ecosystem.^[16]^[25]

References

[1]
A New Family of Horned Dinosauria, From the Cretaceous
Marsh, O. C. (1888). A new family of horned Dinosauria, from the Cretaceous. American Journal of Science, s3-36(216), 477–478.
[2]
https://doi.org/10.1371/journal.pone.0154218
[3]
[PDF] The Ceratopsia
DESCRIPTION BY COPE. Cope's original description of the present genus and species is as follows: #. During the present season F. B. Meek, of Dr. F. V. ...
[4]
Age, Correlation, and Lithostratigraphic Revision of the Upper ...
Introduction. The Upper Cretaceous (Campanian) Judith River Formation is widely exposed in Montana and constitutes the distal part of one of the major eastward ...Introduction · Methods · Formal Changes to Judith... · The Mid-Judith Discontinuity...
[5]
Othniel C. Marsh
Legend has it that the feud between the men began when Marsh paid some of Cope's hired diggers to send fossils to him and not to Cope. Matters became worse in ...Missing: rivalry | Show results with:rivalry
[6]
https://doi.org/10.1080/02724634.1999.10011182
[7]
[PDF] Basal Ceratopsia
A prominent elliptical antorbital fossa with a small antorbital fenestra exists in most basal neo- ceratopsians, except in Archaeoceratops, in which the ...Missing: morphology | Show results with:morphology
[8]
A revision of the Ceratopsia or horned dinosaurs
Charles W. Gilmore of the United States National Museum, while working largely on other forms of bygone life, has also added some notable ceratopsians to ...
[9]
[PDF] MORPHOMETRIC ANALYSIS OF EVOLUTIONARY TRENDS IN ...
ABSTRACT—The cranial anatomy of ceratopsian (“horned”) dinosaurs is well understood, but the postcranial skeleton has been largely ignored in previous ...
[10]
[PDF] Postcranial anatomy of Yinlong downsi \(Dinosauria
Here we provide the first detailed postcranial description of Yinlong downsi based on the holotype and eight other well-preserved skeletons. Yinlong downsi from ...
[11]
https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.24099
[12]
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0126464
[13]
https://www.sciencedirect.com/science/article/pii/S0195667123002665
[14]
Herbivore teeth and diet - The Australian Museum
Ferns, horsetails, club-mosses, conifers, cycads and ginkgoes dominated Triassic and Jurassic menus. The Cretaceous saw an expansion of options with flowering ...On This Page · Which Dinosaurs Didn't Chew... · Which Dinosaurs Chewed Their...Missing: Ceratops browsing angiosperms
[15]
How Triceratops got its face: An update on the functional evolution of ...
Mar 8, 2023 · Enlarged fenestrae can usually be seen bilaterally in dorsal view of the skull on either side of the parietal bar. Exceptions to this are seen ...HORNS AND HORN USE · CRANIAL MUSCULATURE · FEEDING BIOMECHANICSMissing: primitive | Show results with:primitive
[16]
Jaw Mechanics in Basal Ceratopsia (Ornithischia, Dinosauria)
Aug 26, 2009 · Bite force never exceeds the maximum total input force in any of the basal ceratopsians analyzed, with the largest ratio of 0.84 relative to the ...
[17]
Phylogenetic patterns of enamel microstructure in dinosaur teeth
Aug 6, 2025 · The teeth of most early ornithischian dinosaurs were relatively simple triangular or leaf-shaped teeth for shearing plant material [32], but ...
[18]
Dental microwear reveals mammal-like chewing in the ... - NIH
Jul 6, 2016 · This initial inclined palinal direction has also been demonstrated in the basal ceratopsian Psittacosaurus. Previously, Sereno (1987) and ...
[19]
Ontogenetic stages of ceratopsian dinosaur Psittacosaurus in bone ...
Based on a histological study of 43 thin sections from 17 individuals of this species, we recognize four histological ontogenetic stages, i.e., hatchling, ...
[20]
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, ...
[21]
Re-Evaluating Sexual Dimorphism in Protoceratops andrewsi ...
May 7, 2015 · Here, we reassess potential sexual dimorphism in skulls of Protoceratops andrewsi by applying two-dimensional geometric morphometrics to 29 skulls in lateral ...
[22]
Filling in Gaps in the Ceratopsid Histologic Database: Histology of ...
Feb 17, 2019 · Bone histology grants substantial insight into the growth and biology of fossil vertebrates. Many of the major non-avian dinosaurian clades ...
[23]
Transitional evolutionary forms in chasmosaurine ceratopsid ... - PeerJ
Jun 5, 2020 · The parietal fenestrae of ceratopsian dinosaurs open and expand in size through ontogeny ... If so, this would be a paedomorphic trend as ...
[24]
[PDF] JUDITH RIYER BEDS - USGS Publications Warehouse
Ceratops Marsh, 1888, Am. Jour. Sci., 3d ser., vol. 36, p. 477, PI. II ... Fossil plants found in Judith River formation. Fern, indeterminable. Fern ...<|control11|><|separator|>
[25]
Stratigraphy and sedimentology of vertebrate microfossil sites in the ...
A sedimentologic study of 24 vertebrate microfossil sites in the uppermost exposures of the Judith River Formation in and around Dinosaur Provincial Park, ...
[26]
Spiclypeus shipporum gen. et sp. nov., a Boldly Audacious New ...
This study reports on a new ceratopsid, Spiclypeus shipporum gen et sp. nov., from the lower Coal Ridge Member of the Judith River Formation in Montana, USA.<|control11|><|separator|>
[27]
Untitled Document - Paleofile.com
USNM 4802: Squamosal. TREXLER, 1995. Locality: Along the Milk River, Montana. Horizon: Judith River Formation. Biostratigraphy: Age: Judithian age, Campanian ...
[28]
North American nonmarine climates and vegetation during the Late ...
Aug 6, 2025 · Northward of approximately paleolatitude 65°N, evergreen vegetation was replaced by predominantly deciduous vegetation. This replacement is ...
[29]
The “Judith River–Belly River problem” revisited (Montana-Alberta ...
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Fluvial processes and vertebrate taphonomy: the upper cretaceous ...
A fluvial, meandering-channel depositional model has been developed for the upper 60 m of the Judith River sequence in the south-central area of the Park.<|control11|><|separator|>
[31]
Origins of microfossil bonebeds: insights from the Upper Cretaceous ...
Apr 8, 2016 · We studied the sedimentology and taphonomy of 27 microfossil bonebeds in the Campanian Judith River Formation of Montana in order to reconstruct ...Missing: vegetation | Show results with:vegetation<|separator|>
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[PDF] THE VERTEBRATEFAUNA OF THE - AMNH Library Digital Repository
HISTORICAL REVIEW. Stratigraphy of the Judith River Formation. Fauna of the Judith River Formation. ... Dinosaur Egg Shell Fragments.<|control11|><|separator|>
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A transitional species of Daspletosaurus Russell, 1970 from ... - PeerJ
Nov 25, 2022 · A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. Research Article.
[34]
Paleooecology of the Judith River Formation (Campanian) of ...
A survey of the stratigraphic disriubtion of 34 taxa of vertebrates in Judith River Formation (Campanian) of Dinosaur Provincial Park, Alberta, Canada