taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562773	https://doi.org/10.3897/fr.555.e178152.figure2	Figure 2. Block KDRC - SA-V 001 bearing the holotype of Doolysaurus huhmini gen. et sp. nov. A, B. Photographs of KDRC - SA-V 001; C. Digitally rendered models of the bones of KDRC - SA-V 001 from CT scan data. Abbreviations: bo: basioccipital; bs: basisphenoid; co: coracoid; de: dentary; eo: exoccipital-opisthotic; fe: femur; fi: fibula; ga: gastrolith; hu: humerus; mt: metatarsal; na: neural arch; pd: pedal digits; qa: quadrate; ri: rib; sc: scapula; so: supraoccipital; ti: tibia; vc: vertebral centrum.	Figure 2. Block KDRC - SA-V 001 bearing the holotype of Doolysaurus huhmini gen. et sp. nov. A, B. Photographs of KDRC - SA-V 001; C. Digitally rendered models of the bones of KDRC - SA-V 001 from CT scan data. Abbreviations: bo: basioccipital; bs: basisphenoid; co: coracoid; de: dentary; eo: exoccipital-opisthotic; fe: femur; fi: fibula; ga: gastrolith; hu: humerus; mt: metatarsal; na: neural arch; pd: pedal digits; qa: quadrate; ri: rib; sc: scapula; so: supraoccipital; ti: tibia; vc: vertebral centrum.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562774	https://doi.org/10.3897/fr.555.e178152.figure3	Figure 3. Skeletal anatomy of Doolysaurus huhmini gen. et sp. nov. All scale bars are 10 mm. Artwork by Janet Cañamar. Abbreviations: 4 th tr: fourth trochanter; be: buccal emargination; boc: basioccipital condyle; bpro: boss for articulation with proatlas; bt: basal tubera; cc: cnemial condyle; cfo: carotid aorta foramen; cpc: coronoid process; dh: dorsal head; dtt: dentary tooth; ec: endocranial cavity; eoas: exoccipital articular surface; fm: foramen magnum; lc: lateral condyle; lw: lateral wing; mc: medial condyle; mkc: Meckelian canal; mt: metatarsal; mw: medial wing; mxt: maxillary tooth; ns: neural spine; oc: occipital condyle; pd: pedal digit; plp: posterolateral process; pop: paroccipital process; poz: postzygapophysis; prz: prezygapophyses; pscf: posterior semicircular canal foramen; ptf: posttemporal foramen.	Figure 3. Skeletal anatomy of Doolysaurus huhmini gen. et sp. nov. All scale bars are 10 mm. Artwork by Janet Cañamar. Abbreviations: 4 th tr: fourth trochanter; be: buccal emargination; boc: basioccipital condyle; bpro: boss for articulation with proatlas; bt: basal tubera; cc: cnemial condyle; cfo: carotid aorta foramen; cpc: coronoid process; dh: dorsal head; dtt: dentary tooth; ec: endocranial cavity; eoas: exoccipital articular surface; fm: foramen magnum; lc: lateral condyle; lw: lateral wing; mc: medial condyle; mkc: Meckelian canal; mt: metatarsal; mw: medial wing; mxt: maxillary tooth; ns: neural spine; oc: occipital condyle; pd: pedal digit; plp: posterolateral process; pop: paroccipital process; poz: postzygapophysis; prz: prezygapophyses; pscf: posterior semicircular canal foramen; ptf: posttemporal foramen.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562775	https://doi.org/10.3897/fr.555.e178152.figure4	Figure 4. Comparison of the distal femoral morphology in early-diverging neornithischians. A. Distal femur of Koreanosaurus boseongensis in medial view (left), cross-sectional outline at the distal condyle (right), and ventral view of the distal condyle (bottom); B. Distal femur of Doolysaurus huhmini in medial view (left), cross-sectional CT-scanned image (right), and ventral view of the distal condyle (bottom); C. Comparative distal region of other early-diverging neornithischians, including Jeholosaurus (Han et al. 2012), Haya (Barta and Norell 2021), Thescelosaurus assiniboiensis (Brown et al. 2011), and Parksosaurus (Sues et al. 2023). All inset outlines are modified from images in Brown et al. (2011), Han et al. (2012), Barta and Norell (2021), and Sues et al. (2023) and were mirrored to facilitate comparison. All models and drawings are not to scale. Abbreviations: mc: medial condyle; lc: lateral condyle.	Figure 4. Comparison of the distal femoral morphology in early-diverging neornithischians. A. Distal femur of Koreanosaurus boseongensis in medial view (left), cross-sectional outline at the distal condyle (right), and ventral view of the distal condyle (bottom); B. Distal femur of Doolysaurus huhmini in medial view (left), cross-sectional CT-scanned image (right), and ventral view of the distal condyle (bottom); C. Comparative distal region of other early-diverging neornithischians, including Jeholosaurus (Han et al. 2012), Haya (Barta and Norell 2021), Thescelosaurus assiniboiensis (Brown et al. 2011), and Parksosaurus (Sues et al. 2023). All inset outlines are modified from images in Brown et al. (2011), Han et al. (2012), Barta and Norell (2021), and Sues et al. (2023) and were mirrored to facilitate comparison. All models and drawings are not to scale. Abbreviations: mc: medial condyle; lc: lateral condyle.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562776	https://doi.org/10.3897/fr.555.e178152.figure5	Figure 5. Comparison of the proximal tibial morphology in early-diverging neornithischians. A. Proximal tibia of Koreanosaurus boseongensis in lateral view (left), cross-sectional outline at the proximal condyle (right), and dorsal view of the proximal condyle (bottom); B. Proximal tibia of Doolysaurus huhmini in lateral view (left), cross-sectional CT-scanned image (right), and dorsal view of the proximal condyle (bottom); C. Comparative proximal region of other early-diverging neornithischians, including Albertadromeus (Brown et al. 2013), Orodromeus (Scheetz 1999), Thescelosaurus assiniboiensis (Brown et al. 2011), and Parksosaurus (Sues et al. 2023). All inset outlines are modified from images in Scheetz (1999), Brown et al. (2011), Brown et al. (2013), and Sues et al. (2023) and were mirrored to facilitate comparison. All models and drawings are not to scale. Abbreviations: ac: accessory condyle; cc: cnemial condyle; lc: lateral condyle; mc: medial condyle.	Figure 5. Comparison of the proximal tibial morphology in early-diverging neornithischians. A. Proximal tibia of Koreanosaurus boseongensis in lateral view (left), cross-sectional outline at the proximal condyle (right), and dorsal view of the proximal condyle (bottom); B. Proximal tibia of Doolysaurus huhmini in lateral view (left), cross-sectional CT-scanned image (right), and dorsal view of the proximal condyle (bottom); C. Comparative proximal region of other early-diverging neornithischians, including Albertadromeus (Brown et al. 2013), Orodromeus (Scheetz 1999), Thescelosaurus assiniboiensis (Brown et al. 2011), and Parksosaurus (Sues et al. 2023). All inset outlines are modified from images in Scheetz (1999), Brown et al. (2011), Brown et al. (2013), and Sues et al. (2023) and were mirrored to facilitate comparison. All models and drawings are not to scale. Abbreviations: ac: accessory condyle; cc: cnemial condyle; lc: lateral condyle; mc: medial condyle.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562777	https://doi.org/10.3897/fr.555.e178152.figure6	Figure 6. Left maxilla (A) and dentary (B) of the holotype specimen (KDRC - SA-V 001). Abbreviations: ap: anterior process; be: buccal emargination; cpc: coronoid process; ef: elliptical foramen; mkc: Meckelian canal; plp: posterolateral process; rt: replacing tooth.	Figure 6. Left maxilla (A) and dentary (B) of the holotype specimen (KDRC - SA-V 001). Abbreviations: ap: anterior process; be: buccal emargination; cpc: coronoid process; ef: elliptical foramen; mkc: Meckelian canal; plp: posterolateral process; rt: replacing tooth.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562778	https://doi.org/10.3897/fr.555.e178152.figure7	Figure 7. Left quadrate from the holotype specimen (KDRC - SA-V 001). Abbreviations: dh: dorsal head; lc: lateral condyle; lw: lateral wing; mc: medial condyle; mw: medial wing; mwf: medial wing fossa.	Figure 7. Left quadrate from the holotype specimen (KDRC - SA-V 001). Abbreviations: dh: dorsal head; lc: lateral condyle; lw: lateral wing; mc: medial condyle; mw: medial wing; mwf: medial wing fossa.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562779	https://doi.org/10.3897/fr.555.e178152.figure8	Figure 8. Basicranial region of the holotype specimen (KDRC - SA-V 001): A. Left fused exoccipital / opisthotic; B. Supraoccipital; C. Basioccipital; and D. Basisphenoid. Abbreviations: bk: basioccipital keel; boc: basioccipital condyle; bpro: boss for articulation with proatlas; bass: basisphenoid articular surface; bt: basal tubera; cfo: carotid aorta foramen; cri: crista interfenestralis; ec: endocranial cavity; eoas: exoccipital articular surface; fm: foramen magnum; lsas: laterosphenoid articular surface; n: notch; oc: occipital condyle; paas: parietal articular surface; pop: paroccipital process; pras: prootic articular surface; pscf: posterior semicircular canal foramen; ptf: posttemporal foramen; rst: recessus scalae tympani; st: sella turcica.	Figure 8. Basicranial region of the holotype specimen (KDRC - SA-V 001): A. Left fused exoccipital / opisthotic; B. Supraoccipital; C. Basioccipital; and D. Basisphenoid. Abbreviations: bk: basioccipital keel; boc: basioccipital condyle; bpro: boss for articulation with proatlas; bass: basisphenoid articular surface; bt: basal tubera; cfo: carotid aorta foramen; cri: crista interfenestralis; ec: endocranial cavity; eoas: exoccipital articular surface; fm: foramen magnum; lsas: laterosphenoid articular surface; n: notch; oc: occipital condyle; paas: parietal articular surface; pop: paroccipital process; pras: prootic articular surface; pscf: posterior semicircular canal foramen; ptf: posttemporal foramen; rst: recessus scalae tympani; st: sella turcica.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562780	https://doi.org/10.3897/fr.555.e178152.figure9	Figure 9. Dorsal vertebrae and ribs of the holotype specimen (KDRC - SA-V 001): A. Dorsal neural arches; B. Vertebral centra; C. Dorsal ribs. Abbreviations: na: neural arch; ns: neural spine; poz: postzygapophysis; prz: prezygapophyses; tp: transverse process; vk: ventral keel.	Figure 9. Dorsal vertebrae and ribs of the holotype specimen (KDRC - SA-V 001): A. Dorsal neural arches; B. Vertebral centra; C. Dorsal ribs. Abbreviations: na: neural arch; ns: neural spine; poz: postzygapophysis; prz: prezygapophyses; tp: transverse process; vk: ventral keel.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562781	https://doi.org/10.3897/fr.555.e178152.figure10	Figure 10. Pectoral girdle and forelimb elements of the holotype specimen (KDRC - SA-V 001): A. Distal portion of the humerus; B. Possible fragment of the scapula; C. Possible fragment of the coracoid. Abbreviations: rc: radial condyle; uc: ulnar condyle.	Figure 10. Pectoral girdle and forelimb elements of the holotype specimen (KDRC - SA-V 001): A. Distal portion of the humerus; B. Possible fragment of the scapula; C. Possible fragment of the coracoid. Abbreviations: rc: radial condyle; uc: ulnar condyle.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562782	https://doi.org/10.3897/fr.555.e178152.figure11	Figure 11. Left femur of the holotype specimen (KDRC - SA-V 001). Abbreviations: 4 th tr: fourth trochanter; btd: basitrochanteric depression; lc: lateral condyle; mc: medial condyle; pig: posterior intercondylar groove.	Figure 11. Left femur of the holotype specimen (KDRC - SA-V 001). Abbreviations: 4 th tr: fourth trochanter; btd: basitrochanteric depression; lc: lateral condyle; mc: medial condyle; pig: posterior intercondylar groove.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562783	https://doi.org/10.3897/fr.555.e178152.figure12	Figure 12. Left A. Tibia and B. Fibula of KDRC - SA-V 001. Abbreviations: cc: cnemial condyle; lc: lateral condyle; mc: medial condyle.	Figure 12. Left A. Tibia and B. Fibula of KDRC - SA-V 001. Abbreviations: cc: cnemial condyle; lc: lateral condyle; mc: medial condyle.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562784	https://doi.org/10.3897/fr.555.e178152.figure13	Figure 13. Left pes of KDRC - SA-V 001. Abbreviations: af: articular facet; cp: collateral pit; dt: distal trochlea; ep: extensor pit; epc: extensor process; lg: longitudinal groove; mr: medial ridge; mt: metatarsal; pd: pedal digit.	Figure 13. Left pes of KDRC - SA-V 001. Abbreviations: af: articular facet; cp: collateral pit; dt: distal trochlea; ep: extensor pit; epc: extensor process; lg: longitudinal groove; mr: medial ridge; mt: metatarsal; pd: pedal digit.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562785	https://doi.org/10.3897/fr.555.e178152.figure14	Figure 14. Gastroliths of KDRC - SA-V 001. A. Exposed gastrolith mass of KDRC - SA-V 001; B. CT-scanned image of the gastroliths; C. Relationship between body mass and gastrolith mass in dinosaurs, including extant birds. Dotted lines indicate the regression line for each group with more than three samples, and the black line represents the regression line for the total dataset. Gastrolith and body mass data follow the dataset compiled by Wings and Sander (2007), Cerda (2008), Makovicky et al. (2011), Liu et al. (2023), and Chinzorig et al. (2025). The gastrolith mass of Doolysaurus is a minimum value (see text), and the estimated body mass is based on femoral circumference using the equation of Campione et al. (2014). Abbreviations: ga: gastrolith; na: neural arch; vc: vertebral centrum.	Figure 14. Gastroliths of KDRC - SA-V 001. A. Exposed gastrolith mass of KDRC - SA-V 001; B. CT-scanned image of the gastroliths; C. Relationship between body mass and gastrolith mass in dinosaurs, including extant birds. Dotted lines indicate the regression line for each group with more than three samples, and the black line represents the regression line for the total dataset. Gastrolith and body mass data follow the dataset compiled by Wings and Sander (2007), Cerda (2008), Makovicky et al. (2011), Liu et al. (2023), and Chinzorig et al. (2025). The gastrolith mass of Doolysaurus is a minimum value (see text), and the estimated body mass is based on femoral circumference using the equation of Campione et al. (2014). Abbreviations: ga: gastrolith; na: neural arch; vc: vertebral centrum.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562786	https://doi.org/10.3897/fr.555.e178152.figure15	Figure 15. Histological thin-section images of the femoral cortex of KDRC - SA-V 001. A. Cross-section of the anterolateral region of the femur; B, C. Magnified view of the cortical bone under plane-polarized light (B) and cross-polarized light with mineral oil coating (C). Abbreviations: FLB: fibrolamellar bone; LAG: line of arrested growth.	Figure 15. Histological thin-section images of the femoral cortex of KDRC - SA-V 001. A. Cross-section of the anterolateral region of the femur; B, C. Magnified view of the cortical bone under plane-polarized light (B) and cross-polarized light with mineral oil coating (C). Abbreviations: FLB: fibrolamellar bone; LAG: line of arrested growth.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
8DC59F424EDB55289371351242EC2B5B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1562787	https://doi.org/10.3897/fr.555.e178152.figure16	Figure 16. Phylogenetic tree of Thescelosauridae from the tip-dated Bayesian analysis of the character matrix of Fonseca et al. (2024), showing the position of Doolysaurus and scaled to geological time. Small numbers at nodes are posterior probabilities. The color of each tip node corresponds to the continent where the fossil was discovered. Taxa with documented gastroliths (Changmiania, Haya, and Doolysaurus) are highlighted with silhouettes. Tip ages were constrained following the mean of the minimum and maximum ages from the Paleobiology Database (PBDB; http://paleobiodb.org) and Avrahami et al. (2024), with the tip age of Doolysaurus huhmini fixed at 103.45 Ma, corresponding to the midpoint between the Albian and Cenomanian. Doolysaurus huhmini is recovered within Thescelosaurinae, forming a clade with Fona, Parksosaurus, and Thescelosaurus.	Figure 16. Phylogenetic tree of Thescelosauridae from the tip-dated Bayesian analysis of the character matrix of Fonseca et al. (2024), showing the position of Doolysaurus and scaled to geological time. Small numbers at nodes are posterior probabilities. The color of each tip node corresponds to the continent where the fossil was discovered. Taxa with documented gastroliths (Changmiania, Haya, and Doolysaurus) are highlighted with silhouettes. Tip ages were constrained following the mean of the minimum and maximum ages from the Paleobiology Database (PBDB; http://paleobiodb.org) and Avrahami et al. (2024), with the tip age of Doolysaurus huhmini fixed at 103.45 Ma, corresponding to the midpoint between the Albian and Cenomanian. Doolysaurus huhmini is recovered within Thescelosaurinae, forming a clade with Fona, Parksosaurus, and Thescelosaurus.	2026-03-19	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.		Zenodo	biologists	Jung, Jongyun;Kim, Minguk;Jo, Hyemin;Clarke, Julia A.			
