Ecology

Abstract

Ceratosaurs (Ceratosauria), including Ceratosaurus and derived abelisaurids such as Carnotaurus, were the dominant large-bodied predators in Cretaceous ecosystems of the Southern Hemisphere, paralleling the ecological role of tyrannosaurids in the Cretaceous Northern Hemisphere. They are known for their short, deep skulls and prominent cranial ornamentation. These traits have often been interpreted as potential adaptations for intraspecific combat, sexual display, and as a buttress during feeding-induced mechanical stresses. While these adaptations have often been discussed, they have been subject to little mechanical testing, particularly using 3D models due to previous limitations in software. Here we used a mixture of computed tomography (CT) scanning and surface scanning to create accurate 3D models of four ceratosaurian skulls: Ceratosaurus, Masiakasaurus, Carnotaurus, and Majungasaurus. Using finite element analysis, we assessed the mechanical performance of the skull during feeding, notably including the small-bodied noasaurid Masiakasaurus. Our results show that despite their compact shape, large abelisaurs do not exhibit low stress under bite loading. Cranial ornamentation also fails to act as a structural buttress. Additionally, functional analyses of Masiakasaurus combined with its procumbent teeth imply that it was suited for grasping small prey, suggesting niche partitioning with the large sympatric abelisaur Majungasaurus. These findings challenge prevailing assumptions about ceratosaur skull strength and suggest that large abelisaurs fulfilled ecological roles comparable to large tyrannosaurids.

IntroductionCeratosauria is a taxonomically and morphologically diverse clade of basal, non-tetanuran theropod dinosaurs distributed across the Late Jurassic and Cretaceous1. Although theropods are broadly known for achieving large body sizes and for their ecological dominance as bipedal carnivores2,3,4,5, the ceratosaurs represent an under-researched evolutionary pathway within this group. Basal ceratosaurs, such as the Kimmeridgian Late Jurassic Elaphrosaurus, were relatively lightly built6, but by the Maastrichtian Late Cretaceous the group produced large-bodied predators such as Carnotaurus sastrei, which may have exceeded 1,700 kg in body mass7,8,9. The group is commonly associated with the Southern Hemisphere10,11,12, in contrast to tyrannosaurids which were the most prevalent carnivorous dinosaurs of the Cretaceous Northern Hemisphere13,14.Unlike many other large theropods, ceratosaurs evolved a combination of unusual cranial features, including rugose ornamentation, dorsoventrally deepened snouts, and in some taxa, extremely short skulls12. Carnotaurus is commonly noted for its prominent cranial horns15, and Ceratosaurus similarly possessed distinctive horns above the eyes and a midline nasal crest16. Majungasaurus possessed tall, rugose nasals, struts within sinuses, and a horn-like projection of the frontal17. Various explanations have been offered for the functional significance of these traits, including intraspecific combat15,18 and species recognition19.A notable neoceratosaur is Masiakasaurus knopfleri, a small-bodied noasaur from the Maastrichtian of Madagascar with a body mass under 50 kg11,20. Despite its relatively small size, Masiakasaurus possessed a highly modified skull and dentition, including forward-projecting teeth at the anterior of the jaw, possibly indicating specialized feeding behaviors and biomechanics21. The inclusion of Masiakasaurus offers an opportunity to evaluate how skull mechanics and stress distribution vary across a broad range of ceratosaur body sizes and skull morphologies, as well as the biomechanical implications of procumbent teeth in a diapsid.Body size is a key trait affecting feeding mechanics. Larger body size may facilitate higher bite forces due to greater absolute adductor muscle mass but also impose structural constraints that can influence cranial stresses22,23,24. The effects of body size on cranial function remain poorly understood in extinct clades, particularly among ceratosaurs. Large ceratosaurian taxa tend to exhibit deep, robust skulls, whereas smaller taxa display longer, more gracile morphologies (Fig. 1). These size-related changes in skull shape differ from those seen in other large theropod clades: tyrannosaurids reinforce anterior skull regions25, while spinosaurs elongate their rostra26,27, reflecting distinct biomechanical strategies associated with extreme size. Ceratosaurs were also temporally and phylogenetically distinct from other large South American theropods, including Jurassic megalosauroids such as Asfaltovenator28 and Piatnitzkysaurus29, as well as Late Cretaceous megaraptorids, e.g., Joaquinraptor30. This context highlights that the cranial modifications observed in ceratosaurs represent a lineage-specific response to increasing body size rather than a universal pattern among contemporaneous large predators. Fig. 1Simplified Ceratosauria cladogram based on Delcourt12. (A) Carnotaurus; (B) Majungasaurus; (C) Masiakasaurus; (D) Ceratosaurus. 3D models not to scale.Full size imageAdvances in digital modelling techniques, such as surface scanning and 3D finite element analysis (FEA), now allows for the investigation of these questions using 3D reconstructions of fossil skulls (Fig. 2)31,32,33,34. FEA can simulate the mechanical response of a skull to loading, providing estimates of stress and strain as proxies for structural performance. Such studies have increasingly informed our understanding of feeding adaptations in extinct taxa but have yet to focus on ceratosaurs as a lineage.Fig. 2Scanning and FEA pipeline using Masiakasaurus replica skull. (A) Masiaksaurus skull mount at FMNH; (B) surface scan of Masiakasaurus skull in Artec Studio Professional 14; (C) finalized STL file of Masiakasaurus skull in Geomagic Wrap; (D) Masiakasaurus FE model in Abaqus/CAE.Full size imageHere we apply 3D FEA to a series of ceratosaurian taxa spanning a wide range of skull sizes and morphologies: from the gracile, possibly-specialized Masiakasaurus to the deep-skulled, large-bodied Carnotaurus. Our aim is to evaluate how divergent skull morphologies influence cranial stress, mandibular stress, and feeding performance within Ceratosauria using a diverse sample of taxa.We test the following main hypotheses:
Hypothesis 1
Skull stress decreases as ceratosaurs increase in size due to muscle volume.

Hypothesis 2
Skull stress remains constant or increases in large-bodied ceratosaurs due to robust skull architecture in large abelisaurs that permit greater muscle force despite increasing size.
By focusing exclusively on ceratosaurs and incorporating high-resolution digital biomechanical models, this study seeks to provide new insights into how skull and tooth anatomy interact in shaping feeding adaptations in this distinctive clade of predatory dinosaurs.ResultsCranial stress patterns and plotsMajungasaurus experiences the highest cranial stress, followed by Carnotaurus (Figs. 3 and 4). These stresses are most noticeable at the quadrate and quadratojugal due to the positioning of our constraints, though the stresses in Carnotaurus wrap forward onto the jugal. Ceratosaurus similarly experiences higher stresses at the jugal, though overall cranial stresses are not as high as those in the abelisaurs. Masiakasaurus overall experiences the lowest cranial stresses, with the stresses most noticeable at its teeth.Fig. 3Ceratosaur FE results. (A) Carnotaurus; (B) Majungasaurus; (C) Masiakasaurus; (D) Ceratosaurus. Lower skull stresses are indicated by cooler colors, i.e., blue and green. Scale bars represent 25 cm.Full size imageFig. 4Ceratosaur skull lengths plotted against mean von Mises stress for both cranial and mandibular FE models. Silhouette images from PhyloPic by Scott Hartman.Full size imageCranial ornamentation appears to have little effect on absorbing cranial stresses. The horns in Carnotaurus are not mechanically affected by stresses and constraints applied to the quadrate and teeth; likewise, the bony struts of the frontal horn in Majungasaurus are unaffected. The nasal horn in Ceratosaurus appears to offer no functional advantage in feeding; however, the small hornlets above the orbits display moderate stresses.Mandible stress patterns and plotsAs in the cranial results, Majungasaurus experienced the highest mandibular stresses, which are noticeable throughout the FE model (Fig. 3). Similarly, Carnotaurus experienced high mandibular stress, in particular near the anterior end of the dentary. The lower stresses experienced by Masiakasaurus and Ceratosaurus are comparable to each other. Masiakasaurus experienced the lowest overall stresses; the dentary appears less susceptible to breakage than other ceratosaurs in the dataset when forces are applied to the anterior teeth. Overall, there is a clear trend of increasing skull stresses with increasing skull size (Fig. 4), which is particularly notable in the two abelisaurids.
Masiakasaurus biting scenariosThe small-bodied noasaur Masiakasaurus was further tested by applying forces to the midline teeth and posterior teeth to further elucidate the biomechanical implications of procumbent teeth (Fig. 5). We found that when forces were applied only at the front teeth, stresses acting at the mandible were decreased overall, though the quadrate and jugal are more highly stressed. Forces applied at the midline and posterior teeth resulted in much higher stresses acting on the mandible. This may be due to procumbent teeth functioning to isolate or reduce load transfer to the rest of the mandible (see Discussion).Fig. 5Comparison of von Mises stress distributions for different Masiakasaurus biting scenarios. (A) Forces applied to the anterior (procumbent) teeth; (B) forces applied to the midline teeth; (C) forces applied to the posterior teeth. Note the decreased stresses along the mandible in (A).Full size imageDiscussionDespite their distinct cranial morphologies and phylogenetic separation from other theropod lineages, ceratosaurs overall do not appear to exhibit unique biomechanical adaptations for stress resistance during feeding. Cranial ornamentation in these taxa, such as the prominent nasal horn of Ceratosaurus, does not appear to buttress the skull against feeding-induced stress, though the small horns above the orbits, formed by lacrimal bones, do appear stressed during feeding scenarios. Finite element analyses reveal von Mises stress magnitudes in Ceratosaurus that are comparable to similarly sized theropods lacking such ornamentation, suggesting that these cranial features primarily served non-feeding functions, such as species recognition or intraspecific display. The two large abelisaurs in our dataset, Carnotaurus and Majunagasaurus, appear to utilize their short, broad skulls to better absorb high feeding stresses, which is a trait shared by large tyrannosaurids.Our results show that high von Mises stresses in ceratosaur skulls are comparable to those experienced by tyrannosaurids, as the magnitude of stresses increases in large abelisaurids, which has previously been noted in large tyrannosaurids24,35,36,37,38. Previous interpretations of Carnotaurus estimated low bite forces and rapid biting movements15,39. Other interpretations have argued that it possessed a relatively high bite force of ~ 7,000 Newtons40, and at least twice the bite force of the American alligator, which likely possesses the strongest bite of any extant tetrapod41.The parallel evolution of large-bodied ceratosaurs in Gondwana and tyrannosaurids in Laurasia represents a compelling case of functional convergence within non-avian dinosaurs. Despite their disparate cranial morphologies and independent evolutionary histories, both lineages evolved skulls capable of withstanding high feeding stresses as they achieved gigantic size, implying that similar biomechanical constraints shaped apex predation in geographically isolated ecosystems. This pattern aligns with broader examples of convergent ecological expansion in theropods, such as the repeated evolution of deep, reinforced rostra in large-bodied predatory lineages and the niche partitioning documented among large tyrannosaurids and alioramins38,42. Within Ceratosauria itself, previous studies have noted iterative trends toward cranial deepening, reduction of forelimbs, and increasingly specialized feeding morphologies in derived abelisaurids1,43, paralleling structural amplifications seen in later-branching tyrannosaurids. Together with our FE results, these recurring evolutionary trajectories across multiple theropod clades suggest that the demands of subduing large prey frequently channel lineages toward similar biomechanical solutions, even when starting from contrasting anatomical templates.Large abelisaurids including Carnotaurus and Majungasaurus appear to have occupied a biomechanical and ecological space analogous to that of large tyrannosaurids. Like in tyrannosaurids, larger abelisaur skulls appear to be more highly stressed than in ancestral ceratosaur taxa such as Ceratosaurus. Though the ontogeny of Carnotaurus and other large abelisaurs remain relatively unknown, it may be the case as in tyrannosaurids that juvenile forms of derived taxa fed similarly to their adult ancestors37. However, these interpretations should be considered cautiously due to the small sample size of complete abelisaur skulls suitable for 3D FE testing.Cerroni et al.43. argued that the flexibility of Carnotaurus’s mandible may have allowed some degree of kinesis; however, the stiff cranium would prevent flexibility. Our results may reaffirm this work; Carnotaurus and especially Majungasaurus experience the highest mandible stresses, while cranial stresses were noticeably lower, particularly at the anterior of Majungasaurus’s cranium. In general, cranial kinesis is regarded as difficult to prove in any dinosaur species and was likely absent in most44. Additionally, theropod mandible stress magnitudes are almost always higher than their crania due to the size and structural complexity of animal crania compared to mandibles24.Larger ceratosaurs exhibit proportionally higher von Mises stresses than smaller taxa. This pattern aligns with expectations from biomechanical scaling theory. Bite forces are estimated from the summed cross-sectional area of the adductor chamber (PCSA), which scales with the square of linear dimensions, whereas skull cross-sectional area resisting bending scales similarly; however, if skulls were perfectly geometrically similar, stress (force/area) should remain roughly constant across size. Stresses increasing with size indicate that the skulls are not geometrically similar, and that changes in shape modulate how muscle forces are transmitted. Larger abelisaurids such as Carnotaurus and Majungasaurus possess short, deep crania that are likely to improve resistance to dorsoventral bending but also concentrate stresses, while smaller taxa like Ceratosaurus retain elongate skulls that dissipate loads over greater distances. The biomechanical patterns in this study correspond closely with the accelerated evolutionary rates of cranial characters reported for derived abelisaurids45. Their results suggest that the rapid restructuring of the abelisaurid skull, particularly the shift toward short, deep, highly modified facial regions occurred in tandem with the functional changes we document, as these morphologies increasingly shaped how feeding forces were distributed in large-bodied taxa.These findings also bear on previous interpretations of Majungasaurus as a “bite-and-hold” predator17. Such feeding behavior would impose sustained loads on the skull during prey handling and would be facilitated by a stout cranial shape capable of absorbing high stresses without structural failure. Our FE results are consistent with this: Majungasaurus exhibits relatively lower cranial stresses at its anterior than expected given its size, suggesting its skull was mechanically robust under sustained loading, while its mandible shows very high stresses consistent with resisting prey struggling. These findings support the functional plausibility of bite-and-hold feeding in Majungasaurus, though further work on muscle reconstruction and puncture mechanics would be required to test this explicitly. Masiakasaurus is commonly noted for its procumbent anterior dentition, gracile skull, and relatively small body size. Unlike the typical abelisaur skull, which is short and deep, Masiakasaurus’s skull is long and low20. Our FE results support interpretations of a generalist feeding ecology that did not require powerful biting or force transmission and likely focused on small vertebrates and invertebrates, as noted in the related genus Vespersaurus46. Because procumbent teeth project forward, they direct forces more axially along their own long axes, producing lower bending moments at the alveolar margin. More posterior teeth, being upright, transmit bite forces more directly into the mandible, causing higher bending and shear stresses throughout the jaw (Fig. 5). Thus, procumbent teeth are likely specialized for low-stress prey capture or nipping, rather than for generating high bite forces. These functional interpretations also suggest predictions for isolated abelisaurid teeth, which are commonly recovered in Late Cretaceous deposits47. In taxa with gracile, procumbent anterior dentition such as Masiakasaurus, tooth morphology should reflect low bending loads, manifesting as slender crowns with reduced apicobasal wear and minimal chipping. In contrast, the more upright posterior teeth, and those of large-bodied abelisaurids generally, would be expected to show higher frequencies of wear facets, enamel spalling, or microfracturing associated with resisting greater bending and shear forces during prey handling.Given the sympatric relationship between Masiakasaurus and Majungasaurus, it is likely that noasaurs maintained small, slender physiologies into maturity to avoid competition with large abelisaurs. Niche partitioning has been noted in other theropod clades previously, including the alioramins and their larger relatives, e.g., Alioramus and Tarbosaurus38,42. However, the procumbent teeth which are characteristic of Masiakasaurus have not been described in any tyrannosaurid taxa; this may suggest Masiakasaurus utilized these teeth to effectively grasp food items in nutrient or resource-poor environments of Late Cretaceous Madagascar.ConclusionCeratosaurs display a range of biomechanical adaptations to feeding efficiency. As body size increases, abelisaur skulls exhibit higher absolute von Mises stresses, paralleling the pattern observed in tyrannosaurids, despite their independent evolutionary histories and distinct cranial morphologies. This suggests that both lineages experienced similar biomechanical constraints at gigantic size. In large abelisaurs such as Majungasaurus, elevated stresses coincide with cranial shapes that appear well suited to resisting feeding-induced loads, supporting interpretations of relatively high bite forces and possibly a “bite-and-hold” feeding strategy. The small noasaur Masiakasaurus, in contrast, appears to have used its procumbent teeth and low force transmission to capture small vertebrates, invertebrates, or other available resources. Ceratosaurus may have derived minor cranial support from its lacrimal horns, though its prominent nasal horn likely functioned primarily in display. Overall, our results indicate that abelisaurs converged on the biomechanical and ecological roles occupied by tyrannosaurids in the Northern Hemisphere, with both clades evolving skulls capable of tolerating high feeding stresses as they achieved large size.Materials and methodsInstitutional abbreviationsBYUVP: Brigham Young University Museum of Paleontology; DDM: Dinosaur Discovery Museum; FMNH: Field Museum of Natural History; MACN-Pv: Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; UA: University of Antananarivo, Madagascar.Ceratosaur taxonomy and samplingThe earliest ceratosaur representative, Saltriovenator, dates to the Sinemurian Early Jurassic of Italy (199 Ma)48. The clade survived to the Maastrichtian end Cretaceous (66 Ma), where the Madagascan taxa Majungasaurus and Masiakasaurus have been recovered49. Ceratosaurs are generally recognized for their robust, highly ornamented yet short skulls and extremely reduced forelimbs. They are primarily found in the Southern Hemisphere though not exclusively12,50. Ceratosaurus is the most primitive taxon examined in this study and is characterized by its prominent, thin nasal horn which may have functioned for display and/or intraspecific combat51,52,53, as well as unusually fast growth54 and elongated, irregularly formed osteoderms along the midline of its body55.Previous biomechanical studies concerning Carnotaurus have deemed it capable of quick but relatively weak bites, which are critical for capturing small prey15,39. Therrien et al.41. argued that Carnotaurus possessed a bite force twice that of the American alligator, which has one of the highest bite forces in extant species, and it would have been adept at delivering slashing wounds to large prey. Novas18 argued that the well-developed postorbital flanges encircling the orbit in Carnotaurus may have dimmed potential wounds to the eye and head, whereas Cerroni et al.43. assessed the potential flexibility of the skull and determined that any flexibility would be limited to the mandible, as the thickened skull roof and ossification of several cranial joints suggests little or no cranial kinesis.Noasaur feeding biology has remained largely understudied due to lack of skull material. Barbosa et al.46. examined an isolated tooth and two pedal unguals of Vespersaurus paranaensis using 3D FEA and concluded it would have functioned as a generalist feeder preferring small prey and carcasses. This is the first biomechanical study of a 3D noasaur skull. Ceratosaurus, Masiakasaurus, Carnotaurus, and Majungasaurus were included in this study (Fig. 1; Table 1).Table 1 Skull lengths and body mass estimates for each taxon. Skull lengths were measured in MeshLab 2022.02 and masses were referenced from the literature.Full size table3D imagingWe measured skull lengths in MeshLab 2022.02 to better understand possible relationships between size and skull stresses. 3D models originated from both computed tomography (CT) scanning and blue light surface scanning; Carnotaurus is a CT-derived model of the original fossil, while the rest are replicas (Table 1; Table S1).The Masiakasaurus skull used in this study is based on a composite cast housed at FMNH. The original fossil material comprises a partial right maxilla (FMNH PR 2183) and a partial dentary (UA 8680). All remaining cranial and mandibular regions of the cast were reconstructed to restore overall skull shape, based on mirrored copies of preserved elements and comparative morphology from closely related noasaurids described by Sampson et al.20. ; hence, the model is a hypothesis of what a complete, non-deformed specimen would resemble.The Carnotaurus model was segmented from CT data based on the original skull material (MACN-Pv 894). The holotype was CT-scanned using a CT 64 Ingenuity Core medical tomographer, with the skull and left jaw scanned separately. The slice thickness of both scans was about 0.62 mm and scan energy parameters for the skull were 119 mA and 120 kV, and for the lower jaw were 79 mA and 120 kV.Ceratosaurus was surface scanned at the DDM. The DDM cast is based on BYU specimen 12,893, supplemented with comparative material from the larger Ceratosaurus specimens at the Smithsonian National Museum of Natural History and Cleveland Museum of Natural History to complete missing elements; however, the skull material is entirely based on the BYU specimen. The Majungasaurus replica was scanned at the FMNH. It is composed of the cranium with a detached mandible, which made surface scanning easier as the mandible did not obscure the ventral portion of the cranium.Model editingSurface scanned models were created in Artec Studio 14 Professional. The scans were captured at 7–8 frames per second, with the ‘real-time fusion’ option enabled. Crania and mandibles were all scanned separately whenever possible and created as separate 3D object files to avoid large file sizes, since these can cause software slowdown or crashes. Surface scans were oriented together by selecting coordinates and then merging into a single object. Stray pixels and frames with maximum error values above 0.3 were deleted. We applied Global Registration to convert all one-frame surfaces to a single coordinate system using information on the mutual position of each surface pair and then selected ‘Sharp Fusion’ to create a polygonal 3D model, which solidifies the captured and processed frames into an STL file. ‘Sharp Fusion’ best preserves fine details of the scans, including small teeth and rugose bone textures which are particularly common in abelisauroids like Majungasaurus. Lastly, we used the small-object filter to clean the model of floating pixels and the fix holes function to fill any open areas, as the model must be watertight for 3D meshing to proceed without errors (Fig. 2).Once all models were finalized in Geomagic Studio 12, the model surface areas, numbers of triangles, elements, and the volume were calculated and recorded for each model (Table S1, Table S2).Finite element analysesAll models were meshed using HyperMesh (Altair) software (https://altair.com/hypermesh). Element size was set to 10 mm, and in the Tetramesh parameters sub-panel, we selected for optimized mesh quality and for the mesh speed to be gradual. Once the model was meshed successfully, we applied the appropriate material properties to the meshed models. The bone properties were assigned based on crocodilian skull bone: Young’s modulus of 15,000 MPa and Poisson’s ratio of 0.2956,57 for each theropod dinosaur given the close evolutionary relationship between crocodilians and dinosaurs. No sutures were modeled; dentine properties were also not modeled, as they contribute minimally to stress distribution in comparative analyses58.Six total constraint nodes were applied to the quadrate of each cranium and the articular of each mandible to prevent rigid body motion of the models during loading. To achieve this, one node at each location was constrained in a single degree of freedom (DOF) along the X, Y, or Z axis, collectively eliminating translation and rotation while still allowing deformation. Three additional constraint nodes were applied to two premaxillary teeth on each side of the cranium and mandible to simulate contact with prey during a bite. For Masiakasaurus, we additionally conducted FE tests in which bite forces were applied separately to the midline, and posterior teeth to assess how bite point location influenced stress distribution across the skull and elucidate the possible impact of procumbent teeth. Once the mesh was imported into Abaqus/CAE (https://www.3ds.com/products/simulia/abaqus/cae), we applied adductor muscle forces to the models to accurately assess the effects of muscle loading on the skull during a feeding simulation. Locations of muscle insertions were approximated from Holliday59, Rowe & Snively35, and direct study of the specimens. Muscle forces for all theropods were scaled from adult Tyrannosaurus FMNH PR 2081 using the subtemporal fenestra method outlined in Sakamoto60 (Table S3). This was done by measuring the dimensions of the subtemporal fenestra using ImageJ and multiplying the surface area by the isometric muscle tension of 31.5 N/cm2. This method functions as a reliable proxy across amniote clades, and no theropod dinosaur taxa appear to be outliers61.We generated von Mises stress data for each theropod dinosaur cranial and mandibular model. 3D finite element stress maps were generated for each 3D model analyzed, with von Mises stress limits ranging from 0 to 25 MegaPascals in the display figures. Mesh-weighted arithmetic mean (MWAM) values were calculated using RStudio for each theropod specimen to reduce possible discrepancies between CT-derived models and surface scan-derived models and account for element volume differences between models62,63,64. Rowe & Rayfield34 concluded that 3D FE models with a high degree of editing applied prior to meshing tend to output relatively high von Mises stress and maximum principal strain relative to un-modified models. Mesh-weighted von Mises stress data was compiled and compared for all skulls in a phylogenetic context to visualize the effects of large skulls on feeding-induced stresses (Fig. 3). Given that our dataset includes only four taxa, formal phylogenetic generalized least squares (PGLS) regression was not attempted, as such analyzes are statistically underpowered and would yield unstable estimates of phylogenetic signal. Instead, we present von Mises stress values quantitatively and descriptively within a phylogenetic framework.

Data availability

FE files available upon request from the corresponding author, Andre J. Rowe ([email protected]).
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Download referencesFundingWe thank Nick Wiersum for access and assistance with scanning the Ceratosaurus at the DDM. Thanks to William Simpson for access to specimens at the FMNH which allowed us to scan Masiakasaurus and Majungasaurus skull material. Thanks to Martin Ezcurra, Belen von Baczko, and Agustin Martinelli for allowing access to the Carnotaurus holotype and to Salguero Diagnostic Center (Buenos Aires, Argentina) for the CT scanning of the specimen. E.J.R. was funded by BBSRC grant BB/W00867X/1.Author informationAuthors and AffiliationsSchool of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UKAndre J. Rowe & Emily J. RayfieldConsejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Buenos Aires (CABA), ArgentinaMauricio A. CerroniÁrea Laboratorio e Investigación, Museo Municipal “Ernesto Bachmann” (MEB), Dr. Natali S/N 8311, Villa El Chocón, Neuquén, ArgentinaMauricio A. CerroniLaboratorio de Anatomía Comparada y Evolución de los Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, ArgentinaMauricio A. CerroniAuthorsAndre J. RoweView author publicationsSearch author on:PubMed Google ScholarMauricio A. CerroniView author publicationsSearch author on:PubMed Google ScholarEmily J. RayfieldView author publicationsSearch author on:PubMed Google ScholarContributionsA.J.R and E.J.R. conceived of the research. A.J.R. and M.A.C. performed the research. A.J.R., M.A.C., and E.J.R. wrote the manuscript.Corresponding authorCorrespondence to
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Reprints and permissionsAbout this articleCite this articleRowe, A.J., Cerroni, M.A. & Rayfield, E.J. Southern hemisphere ceratosaurs evolved feeding mechanics paralleling those of Northern hemisphere tyrannosaurids.
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Abstract

Persistent organic pollutants (POPs), such as the legacy contaminants polychlorinated biphenyls (PCBs) and dichlorodiphenyltrichloroethane (DDTs), as well as trace elements (TEs) pose a significant risk to marine ecosystems due to their toxicity, persistence, and bioaccumulative nature. Despite regulatory bans, PCBs and DDTs continue to be detected in the marine environment, while TEs levels remain conspicuous as a result of both natural and anthropogenic sources. In this study, we investigated the presence, concentrations, and spatial distribution of 32 PCB congeners, 6 DDT compounds, and 16 TEs in zooplankton collected from 40 sites across the Northwestern Mediterranean Sea. Results revealed widespread contamination, with PCBs detected in all samples (53.2 ± 63.0 ng g-1 dw) and DDTs present in over half the samples (5.1 ± 6.0 ng g-1 dw). Hotspots of POPs contamination were identified near Marseille, and in the wider Gulf of Lion, Barcelona, and the Ebro River mouth. All TEs were detected in zooplankton, with high concentrations of essential TEs, but also elevated levels of toxic elements such as Hg, Pb, and Cd in certain locations such as Barcelona, the Gulf of Lion, Balearic Islands, and southwestern Corsica. Spatial patterns of contamination were strongly linked to urban, industrial, riverine, and historical mining inputs. These results underline the important role of zooplankton as bioindicators for assessing pollutant transfer at the base of the marine food web. They also highlight the urgent need for integrated, long-term monitoring strategies to better understand contaminant dynamics and mitigate ecological risks in the Mediterranean Sea.

Data availability

The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.
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Reprints and permissionsAbout this articleCite this articleBoldrocchi, G., Villa, B., Banfi, D. et al. Presence, levels, and distribution of organic and elemental pollutants in Zooplankton from the Northwestern Mediterranean sea.
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AbstractUrban planners increasingly recognize the importance of improving access to nature to support human well-being, social inclusion, and urban sustainability. However, assessing accessibility to urban green spaces remains challenging, particularly in rapidly urbanizing cities of the Global South, where mobility constraints and socio-spatial inequalities shape who can effectively benefit from these spaces. Urban wetlands, despite providing multiple ecosystem services, are rarely incorporated into formal green infrastructure networks and are often excluded from accessibility assessments. This study examines how integrating urban wetlands as Nature-Based Solutions (NbS) modifies patterns of access to nature in Concepción, Chile. We combine land-use data on parks and wetlands with accessibility measures and origin–destination travel survey data to model walking accessibility under two scenarios: green spaces excluding wetlands and green spaces including wetlands. A Random Forest model is applied to capture heterogeneity across socio-demographic profiles, including age, gender, employment status, and driver’s license ownership. Results show that incorporating urban wetlands significantly increases the accessible surface of green space across the city, with particularly strong gains for groups with limited mobility options, such as women, older adults, unemployed residents, and individuals without access to private vehicles. In peripheral and underserved areas, wetlands partially compensate for deficits in formal park provision, reducing accessibility gaps. These findings provide empirical evidence that urban wetlands function as grounded NbS by redistributing access to nature and underscore the importance of formally integrating wetlands into urban planning instruments to promote more inclusive and resilient cities.

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Data availability

The data that support the findings of this study are derived from a combination of publicly available spatial datasets and restricted-access origin–destination travel survey data. Public spatial data sources are cited in the manuscript. Access to the origin–destination survey data is subject to data protection and confidentiality agreements and is therefore available from the corresponding author upon reasonable request and with permission from the data provider. Derived datasets and code generated during the analysis are available from the corresponding author upon reasonable request.
Code availability

The data that support the findings of this study are derived from a combination of publicly available spatial datasets and restricted-access origin–destination travel survey data. Public spatial data sources are cited in the manuscript. Access to the origin–destination survey data is subject to data protection and confidentiality agreements and is therefore available from the corresponding author upon reasonable request and with permission from the data provider. Derived datasets and code generated during the analysis are available from the corresponding author upon reasonable request.
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Download referencesAcknowledgements“Accesibilidad y valor percibido de humedales urbanos y parques de humedales: Soluciones basadas en la naturaleza para la sustentabilidad de ciudades costeras” ANID FONDECYT REGULAR Nº 1250128. Fondo de Investigación en Ciencia y Tecnología de la ANID. Centro de Interés Nacional “Center for Sustainable Urban Development” (CEDEUS) (2025-2030). ANID CIN 250009.Author informationAuthors and AffiliationsInstituto de Estudios Urbanos y Territoriales, Providencia, ChileCarolina Rojas QuezadaCentro de Desarrollo Urbano Sustentable (CEDEUS), Pontificia Universidad Católica de Chile, Santiago, ChileCarolina Rojas QuezadaSchool of Earth, Environment, and Society, McMaster University, Hamilton, ON, CanadaAntonio PáezCity Lab Biobío, Concepción, ChileHelen de la FuenteCentro de Desarrollo Integral de los Territorios (CEDIT), Pontificia Universidad Católica de Chile, Santiago, ChileBryan CastilloAuthorsCarolina Rojas QuezadaView author publicationsSearch author on:PubMed Google ScholarAntonio PáezView author publicationsSearch author on:PubMed Google ScholarHelen de la FuenteView author publicationsSearch author on:PubMed Google ScholarBryan CastilloView author publicationsSearch author on:PubMed Google ScholarContributions(C.R.): Conceptualization; Writing – original draft; Methodology design. (A.P.): Methodology design; Writing – review and editing. (H.F.): Data curation; Visualization; Machine learning analysis; Calculations. (B.C.): Machine learning analysis; Calculations.Corresponding authorCorrespondence to
Carolina Rojas Quezada.Ethics declarations

Competing interests
The authors declare no competing interests.

Additional informationPublisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
Reprints and permissionsAbout this articleCite this articleQuezada, C.R., Páez, A., Fuente, H.d.l. et al. Grounding Urban wetlands as nature-based solutions: enhancing accessibility to nature in Concepción, Chile.
npj Urban Sustain (2026). https://doi.org/10.1038/s42949-026-00339-8Download citationReceived: 21 August 2025Accepted: 11 January 2026Published: 21 January 2026DOI: https://doi.org/10.1038/s42949-026-00339-8Share this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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AbstractAs global food systems face mounting sustainability pressures, insects are gaining attention as a promising alternative protein source. Yet, entomophagy remains culturally unfamiliar or stigmatized in many Western countries, including Canada. This study investigates attitudes toward insect consumption among 252 adult visitors to the Montreal Insectarium, a public institution promoting insect education and biodiversity awareness. Participants completed a structured questionnaire evaluating willingness to consume various insect-based foods, motivations and barriers, and demographic predictors of acceptance. Overall, 44% of participants reported openness to eating insects (18% had previously consumed them and 26% were willing to try), though fewer were willing to include them in their regular diet (27%) or prepare them at home (17%). Acceptance was highest for products where insect content was less visible, such as baked goods made with insect flour. Key motivators included curiosity, perceived health benefits, and environmental concern, while major deterrents were disgust, food safety concerns, and insect-related fears. Ordinal logistic regression analyses revealed consistent gender effects, with men significantly more willing than women to consume a variety of insect-based foods. Men also showed greater prior experience with insect consumption and were more likely to include insects in their diets or try them in restaurants. Age alone was not a consistent predictor, but significant interactions with gender revealed a complex interplay between these predictors. Moreover, participants with graduate degrees showed greater openness to experimenting with insect-based ingredients when cooking, and prior insect consumption increased with education among women. Overall, our results show that demographic differences (especially gender and education) shape openness to entomophagy more strongly than age alone, suggesting that targeted outreach could be a better strategy than generalized promotion in encouraging insect-based food adoption.

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Data availability

The data used in this study are available from the Figshare Repository: https://doi.org/10.6084/m9.figshare.29429660.
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Rassim Khelifa.Ethics declarations

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Ethical approval
Ethical approval (Certification Number: 30020523) was obtained from Concordia University.

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Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
Reprints and permissionsAbout this articleCite this articleVelchovska, N., Khelifa, R. Acceptance of entomophagy among Canadians at an insectarium.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-35288-wDownload citationReceived: 28 June 2025Accepted: 05 January 2026Published: 21 January 2026DOI: https://doi.org/10.1038/s41598-026-35288-wShare this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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KeywordsInsect consumptionAlternative proteinConsumer attitudesGenderCulinary innovationEnvironmental sustainabilityCanada More

AbstractAccurate and scalable taxonomic classification is essential for biodiversity research, supporting systematic species identification across multiple hierarchical ranks. However, current image-based classification methods often fail to enforce taxonomic consistency, a critical limitation that undermines the reliability of their outputs for scientific use. Additionally, field-based biodiversity studies are constrained by limited computational resources and network availability on edge devices. To address these challenges, this paper proposes TaxonomyNet, an ensemble detection model with six independent heads for taxonomic classification, achieving high detection performance across all ranks (mAP: 90.7–99.75%) after training on a dataset of 50 Australian animal species. Furthermore, to resolve the core challenge of prediction inconsistency, we introduce the Weighted Agreement Loss (WAL) metric–a confidence-weighted disagreement measure designed to enforce structural coherence between predicted outputs and a reference taxonomy. Crucially, the application of this consistency-enforcing mechanism enhances hierarchical classification reliability, improving final species-level accuracy by up to 3.87% compared to baseline and recent published domain-specific foundation models, while also demonstrating superior computational efficiency, reducing delay by 22 minutes across 1500 samples and making it highly suitable for deployment on edge devices. This work provides a practical and extensible solution for reliable hierarchical classification in real-world biodiversity monitoring scenarios.

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Data availability

The dataset of 50 species used in this study is publicly available at https://cutt.ly/gwqHHm8D. The datasets used for the higher taxonomic ranks (genus, family, order, class, and phylum) are derived from the same underlying image collection but differ in naming conventions and label structures. These higher-rank datasets can be provided by the corresponding author upon reasonable request.
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KeywordsImage recognitionYOLODeep learningAustralian speciesConvolutional neural network More

Abstract

Allelopathy is one of the important mechanisms for the spread and expansion of invasive alien plants. The current research mainly focuses on interspecific allelopathy, while there are relatively few studies on intraspecific allelopathy. Solanum rostratum Dunal is an annual invasive plant with strong invasiveness, the secondary metabolites produced by the litter of S. rostratum can accumulate in the soil, and may affect the growth of its own seedlings. Therefore, it is of great significance to clarify the intraspecific allelopathy of S. rostratum for understanding the invasion mechanism or proposing new prevention and control strategies. In this study, the extract of S. rostratum litter was used to treat its seedlings, and the soil physical and chemical properties, soil metabolites, and soil microorganisms were measured to analyze their correlation with the growth of seedlings. The results showed that 0.1 and 1 g L− 1 treatment significantly promoted the leaf area and biomass of seedlings, while 10 g L− 1 treatment significantly inhibited plant height, leaf area index, biomass, net photosynthetic rate, transpiration rate, and stomatal conductance. Some bacteria, such as Brevundimonas alba, Brevundimonas, Altererythribacter, Novosphingobium resinovorum, and Novosphingobium exhibited a higher abundance under 10 g L− 1 treatment, showed a negative correlation with seedling growth. And 25 metabolites detected in the soil, such as 2-Aminobenzoic acid, 2, 6-dibromophenol and palmitaldehyde, could as autotoxins for subsequent functional validation. The results can not only supplement the invasion mechanism of invasive plants from the perspective of intraspecific allelopathy, but also provide theoretical support for formulating control strategies for the S. rostratum.

Data availability

The sequencing data have been deposited in the National Center for Biotechnology Information Sequence Read Archive database (http://www.ncbi.nlm.nih.gov/sra) with BioProject accession number PRJNA1274989.
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Download referencesFundingThis research was supported by a special grant from the Natural Science Foundation of the Xinjiang Uygur Autonomous Region (grant number: 2023D01B37), the National Natural Science Foundation of China (32460684), the Xinjiang Key Laboratory of Soil and Plant Ecological Processes (grant number: 23XJTRZW19) and the Major Science and Technology Public Relations Project Fund of the Science and Technology Department of Xinjiang Uygur Autonomous Region (2023A02006).Author informationAuthor notesThese authors contributed equally: Yuxuan Ma and Lamei Jiang.Authors and AffiliationsXinjiang Key Laboratory for Ecological Adaptation and Evolution of Extreme Environment Organisms, College of Life Sciences, Xinjiang Agricultural University, Ürümqi, 830052, ChinaYuxuan Ma, Lamei Jiang, Shuai Liu, Huixian Liu, Juan Qiu, Shanshan Wang & Dunyan TanCollege of Chemistry, Xinjiang University, Ürümqi, PR ChinaShuai LiuXinjiang Key Laboratory of Soil and Plant Ecological Processes, College of Resources and Environment, Xinjiang Agricultural University, Ürümqi, ChinaGuohao ZhaiState Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Ürümqi, 830011, ChinaDunyan TanAuthorsYuxuan MaView author publicationsSearch author on:PubMed Google ScholarLamei JiangView author publicationsSearch author on:PubMed Google ScholarShuai LiuView author publicationsSearch author on:PubMed Google ScholarHuixian LiuView author publicationsSearch author on:PubMed Google ScholarGuohao ZhaiView author publicationsSearch author on:PubMed Google ScholarJuan QiuView author publicationsSearch author on:PubMed Google ScholarShanshan WangView author publicationsSearch author on:PubMed Google ScholarDunyan TanView author publicationsSearch author on:PubMed Google ScholarContributionsConceptualization, S. Wang and D. Tan; methodology, S. Wang; formal analysis, Y. Ma, H. Liu and L. Jiang; investigation, Y. Ma, L. Jiang, H. Liu and G. Zhai; resources, Y. Ma, and J. Qiu; data curation, Y. Ma, L. Jiang and S. L; writing—original draft preparation and the revision of the article, Y. Ma, L. Jiang, S. W; writing—review and editing, S. Wang and D. Tan; supervision, S. Wang and D. Tan; project administration, S. Wang and D. Tan; funding acquisition, S. Wang and D. Tan. All the authors have read and agreed to the published version of the manuscript.Corresponding authorsCorrespondence to
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AbstractMeeting the ambitious targets set by the Convention on Biological Diversity (CBD) Kunming-Montreal Global Biodiversity Framework (GBF) will require expanding ecosystem restoration across governance domains for marine and coastal ecosystems. Marine spatial planning (MSP), which balances the development of multiple human uses in the ocean with the preservation of ecosystem health, might be the most effective vehicle for achieving this aim. However, to date, MSP and restoration efforts have proceeded on separate tracks, and biodiversity loss continues. In this Perspective we detail how embedding restoration into planning at the site scale, alongside subnational or national MSP, can help to achieve multiple simultaneous global biodiversity targets. We outline four avenues to action that support biodiversity-positive outcomes over the long term: ensure flexible and multi-scaled approaches; make planning community-centred; maintain social–ecological connectivity; and assess climate-related risks and opportunities. We then propose metrics that CBD Parties could use to implement and track progress on integrating ecoscape restoration into climate-smart MSP. With the world’s focus on the GBF and the approaching 2030 deadline, implementing the proposed avenues to action could lead to more rapid achievement of global targets.

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Fig. 1: Examples of ecoscape restoration actions that could be catalysed by climate-smart marine spatial planning, considering interlinked land, riverine and marine areas.Fig. 2: Four key avenues to action for integrating ecoscape restoration into climate-smart marine spatial planning.

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AbstractGreen development of agriculture (GDA) is a critical issue for advancing agricultural modernization and serves as an essential foundation for comprehensive rural revitalization. The impact of GDA on the common prosperity of farmers in rural areas remains controversial. Therefore, this study develops an evaluation index system to measure the level of GDA based on the panel data of prefecture-level cities in China from 2013 to 2022. Subsequently, this paper analyzes the specific effects, mechanisms, regional differences and threshold effects of GDA on common prosperity using the fixed effect model, mediated-effects models, group regression method and threshold effect model. The results show that: Firstly, GDA has a positive impact on the common prosperity for farmers and rural areas. Secondly, the extension of the agricultural industry chain and the expansion of agricultural multi-functionality are important paths for GDA to promote common prosperity. Thirdly, there is regional heterogeneity in the impact of GDA on common prosperity. GDA has a significant impact on common prosperity in eastern China and non-food-producing regions, but it does not lead to common prosperity in western China. Specifically, the GDA in the central region and food-producing areas can only increase farmers’ income and narrow the rural income gap, but it cannot bridge the urban-rural income gap. In addition, the influence of GDA on common prosperity has a threshold effect based on agricultural industrial agglomeration. The common prosperity effect of GDA gradually becomes prominent as the agricultural industrial agglomeration is raised to a certain level.

Data availability

The data presented in this study are available on request from the Corresponding author.
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Download referencesFundingGeneral Projects of the National Social Science Fund(Grant No. 22BGL180); Jiangxi Management Science Decision Consulting Project (20244BAA10026); Henan Province Philosophy and Social Science Planning Youth Project(Grant No. 2023CJJ181).Author informationAuthors and AffiliationsSchool of Economics and Management, Jiangxi Agricultural University, Nanchang, 330000, ChinaZhaoguang Li, Qinying Shi & Kai HuSchool of Economics and Management, Henan Institute of Science and Technology, Xinxiang, 453000, ChinaZhuanqing ChenAuthorsZhaoguang LiView author publicationsSearch author on:PubMed Google ScholarQinying ShiView author publicationsSearch author on:PubMed Google ScholarKai HuView author publicationsSearch author on:PubMed Google ScholarZhuanqing ChenView author publicationsSearch author on:PubMed Google ScholarContributionsConceptualization, Z.L.; methodology and software, K.H.; validation,.;formal analysis, Z.L.; investigation, Z.L.; resources, K.H.; data curation, K.H.; writing—original draft preparation, Z.C.; writing—review and editing,.; visualization, Z.C.; supervision, K.H.; project administration, K.H.; funding acquisition, K.H. All authors have read and agreed to the published version of the manuscript.Corresponding authorsCorrespondence to
Qinying Shi or Kai Hu.Ethics declarations

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Reprints and permissionsAbout this articleCite this articleLi, Z., Shi, Q., Hu, K. et al. The impact of agricultural green development on common prosperity for farmers in rural areas.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-35978-5Download citationReceived: 03 February 2025Accepted: 09 January 2026Published: 20 January 2026DOI: https://doi.org/10.1038/s41598-026-35978-5Share this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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KeywordsGreen development of agricultureCommon prosperityAgricultural industrial chainAgricultural multifunctionalityThreshold effectIncome gap More