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Lesothosaurus

Lesothosaurus diagnosticus is a species of small, basal ornithischian dinosaur known from the Early Jurassic upper Elliot Formation of southern Africa, dating to approximately 200–195 million years ago.^[1] It was a lightweight, bipedal animal with a slender build, grasping hands, and long hind limbs adapted for agile movement, reaching an estimated adult body length of up to 2 meters and a weight of around 20 kilograms based on larger postcranial elements.^[2] The dinosaur possessed a small skull with large eyes, low-crowned teeth featuring coarse denticles suitable for processing soft plant material, and evidence suggests a predominantly herbivorous diet supplemented by occasional animal matter such as insects.^[3] Named by paleontologist Peter Galton in 1978 based on specimens from Lesotho, Lesothosaurus was initially considered a primitive member of the ornithopod group but is now recognized as a basal ornithischian, offering key insights into the early diversification of bird-hipped dinosaurs.^[2] The type specimens, including partial skeletons, represent juveniles, while more recent discoveries from bonebeds in South Africa's Free State province reveal larger individuals and indicate ontogenetic changes in cranial features like increased tooth count and rugosity.^[2] These assemblages, comprising multiple co-occurring individuals, provide the earliest evidence of potential gregarious behavior among ornithischians.^[4] The anatomy of Lesothosaurus, detailed through CT scans and digital reconstructions, highlights primitive traits such as a generalized dentition and postcranial skeleton without specialized defenses, underscoring its role as a generalized early herbivore in a landscape dominated by conifers and ferns.^[5] Its discovery has informed phylogenetic analyses, positioning it near the base of Ornithischia and contributing to debates on the group's Triassic origins and rapid Jurassic radiation.^[6]

Discovery and naming

Discovery

The first fossils attributable to Lesothosaurus were collected in 1959 by a French paleontological team led by François Ellenberger, including Jean Fabre and Léonard Ginsburg, from the Upper Elliot Formation at Likhoele in the Mafeteng district of Lesotho. This material consisted of a right dentary fragment bearing three teeth, later described as the holotype of Fabrosaurus australis by Ginsburg in 1964.^[7] Additional specimens were recovered during subsequent expeditions, including those by University College London in 1963–1964 and a joint effort involving the British Museum of Natural History, Birkbeck College (University of London), Yale University, and the South African Museum in 1967–1968, from sites in Lesotho and the Qacha's Nek area of Lesotho. These efforts yielded over 50 individuals, including disarticulated skeletal elements from multiple localities in the Upper Elliot Formation. The syntypes of Lesothosaurus diagnosticus—BMNH RU B17 (a partial skeleton representing at least two individuals) and BMNH RU B23 (a partial skull)—were among the key finds from these collections, now housed in the Natural History Museum, London. Referred specimens include BMNH R 8501 (a nearly complete skull) and BMNH RU B506 (bonebed material).^[7]^[8] A notable bonebed at the Fouriesburg locality in South Africa's Free State Province, derived from material collected in earlier expeditions but described in 2016, preserved remains of at least three Lesothosaurus individuals, including larger adults, indicating ontogenetic variation and potential gregarious behavior. A 2022 analysis of this bonebed provided the earliest evidence of social living among ornithischians.^[9]^[4] Overall, the known fossil inventory for Lesothosaurus includes over 50 specimens, predominantly juveniles, all derived from the Upper Elliot Formation (Hettangian–Sinemurian stages of the Early Jurassic).^[10]

Naming

The fossils collected from Qacha's Nek in Lesotho, including a partial skeleton (MNHN LES 17), were referred by R. A. Thulborn in 1970 to the existing species Fabrosaurus australis (named by Ginsburg in 1964 based on holotype MNHN LES 9, a dentary fragment) from the Upper Elliot Formation.^[5] In 1978, Peter M. Galton erected the new genus and species Lesothosaurus diagnosticus for the British Museum material, considering Fabrosaurus australis a nomen dubium due to its inadequate diagnosis. The generic name derives from "Lesotho," the country encompassing the type locality, combined with the Greek σαῦρος (sauros), meaning "lizard"; the specific epithet "diagnosticus" refers to the distinctive anatomical features of the type specimens that allow confident referral of additional material. Galton designated the syntypes as the partial skeleton and skull (BMNH RU B17 and BMNH RU B23).^[5] In 2005, a partial skeleton from the upper Elliot Formation in South Africa was described as the new taxon Stormbergia dangershoeki by R. J. Butler (with contributions noted in related works by Knoll et al.). This name was later regarded as a junior subjective synonym of Lesothosaurus diagnosticus in a 2017 study by M. G. Baron, D. B. Norman, and P. M. Barrett, who identified shared autapomorphies such as the loss of metatarsal V and overlapping morphological variation attributable to ontogeny rather than distinct taxa.^[1] Lesothosaurus diagnosticus remains the sole valid species in the genus, with subsequent revisions confirming its monospecific status; no additional species have been recognized since the 2017 synonymy.^[11]

Description

Skull and dentition

The skull of Lesothosaurus diagnosticus is small and elongate, typically measuring 130–150 mm in basal length from the anterior premaxilla to the posterior basioccipital, and appears narrow and triangular in dorsal view. It exhibits primitive ornithischian features, including large antorbital and infratemporal fenestrae that occupy significant portions of the lateral skull surface, as well as sub-rectangular infratemporal fenestrae that extend nearly the full height of the skull. The orbits are proportionally large, comprising approximately 36% of the basal skull length and suggesting enhanced visual capabilities.^[12]^[12] Most known specimens represent juveniles, with larger individuals showing ontogenetic changes such as increased cranial robusticity and tooth count.^[2] The braincase is compact and shows signs of immaturity in known specimens, with unfused elements such as the basisphenoid and basioccipital, alongside elongated, horizontally directed paroccipital processes that project posteriorly from the occiput. These processes contribute to a broad occipital region typical of early ornithischians. The overall cranial architecture, with its prominent fenestrae and arched frontals providing a rounded dorsal profile, underscores Lesothosaurus's basal position within Ornithischia.^[12]^[13] The dentition consists of 12–15 teeth in the maxilla and 12–17 positions in the dentary, though estimates suggest up to 20 dentary teeth in larger individuals; the premaxilla bears five to six smaller, more recurved teeth per side. Tooth crowns are low and leaf-shaped, with asymmetrical profiles featuring a prominent central ridge flanked by coarse marginal denticles, and a distinct cingulum at the base separating the crown from the root. These features indicate a shearing occlusal mechanism, with oblique wear facets on opposing teeth facilitating food processing. Replacement teeth are evident, often positioned lingually to functional ones.^[12]^[12] The jaws include a dentigerous premaxilla forming the rostral tip, paired with a lower jaw that has a spout-shaped, inturned symphysis and a predentary bone contributing to a beak-like structure for cropping vegetation. The straight tooth rows align such that the craniomandibular joint lies slightly below the occlusal plane, and a subtle diastema between tooth rows supports the presence of buccal soft tissues. Analysis of tooth wear patterns reveals double facets consistent with intraoral trituration, providing evidence for fleshy cheeks that enclosed the dentition during feeding; similar wear suggests the possible presence of lips covering the teeth when the jaws were closed.^[12]

Postcrania

The postcranial skeleton of Lesothosaurus diagnosticus is characterized by a gracile, bipedal build adapted for agility, with forelimbs significantly shorter than the hindlimbs. The axial skeleton includes an estimated nine cervical vertebrae, though preserved series are incomplete; these feature low, hook-like neural spines that suggest a flexible neck region. The dorsal series is partially known, with at least five preserved vertebrae exhibiting rectangular neural spines, while the sacral series comprises five vertebrae with trapezoidal to rectangular neural spines and spool-shaped centra bearing reniform articular surfaces.^[1] The pectoral girdle consists of a small, elongate scapula measuring approximately 75–132 mm in length across specimens, with a bowed shaft and expanded distal blade; it is typically 10–15% longer than the humerus and articulates with a coracoid that possesses a prominent foramen. The forelimbs are short and reduced, with the humerus ranging from 72–112 mm in length, featuring a bulbous head, prominent deltopectoral crest extending over 40% of its length, and a reduced olecranon process on the ulna, indicating limited arm mobility. In the pelvic girdle, the ilium is notably elongated and deep, with a laterally compressed preacetabular process and a short postacetabular process, alongside a short prepubis and a pubis that is slightly retroverted. The hindlimbs are robust relative to the forelimbs, dominated by a long femur measuring 103–235 mm, which is bowed and bears a pendant fourth trochanter; the tibia and fibula are slender and subequal in length to the femur, supporting a four-toed pes with the phalangeal formula 2-3-4-3-0 (digit V reduced) and recurved claws on digits I–IV, consistent with cursorial habits.^[1] Overall, L. diagnosticus exhibits hindlimb-dominated proportions, with the humerus comprising about 63% of femur length in smaller specimens, reflecting a bipedal stance; smaller individuals are estimated to have weighed 5–10 kg, while larger adults reached up to around 20 kg, based on femoral dimensions and volumetric models.^[1]^[14]^[15]

Classification

Historical classification

Lesothosaurus diagnosticus was formally named and described by Peter Galton in 1978, based on material previously referred to Fabrosaurus australis by Richard Thulborn in 1970; Galton positioned it as a basal ornithischian within the newly proposed family Fabrosauridae, alongside other small, primitive forms like Fabrosaurus and Alocodon from the Early Jurassic of southern Africa.^[16]^[17] Thulborn's earlier work had interpreted the syntype skulls (NHMUK PV RU B17 and B23) as belonging to a Triassic ornithischian with ornithopod affinities, emphasizing features like the dentition and jaw mechanics that suggested a primitive herbivorous lifestyle.^[16] Galton's classification reinforced this by grouping Lesothosaurus with other "fabrosaurs" as early ornithopods, distinct from more derived ornithischians, based on shared traits such as bipedal posture and leaf-shaped teeth.^[17] In the 1980s and 1990s, interpretations shifted amid debates over its precise affinities, with Thulborn (1971, 1972) and Galton (1972, 1986) viewing Lesothosaurus as a primitive member of either Fabrosauridae or Heterodontosauridae, highlighting similarities in skull morphology and postcranial proportions to South African heterodontosaurids like Heterodontosaurus.^[17] These placements sparked discussions on whether it represented an early ornithopod or a more basal ornithischian, as some features—like the large mandibular fenestra and reduced pubic peduncle—suggested exclusion from the clade Genasauria, a group encompassing more advanced ornithischians such as ceratopsians and ornithopods, as defined by Sereno (1986).^[6] Sereno's detailed redescription in 1991 further argued that Fabrosauridae was a paraphyletic assemblage of basal ornithischians, with Lesothosaurus as one of the most primitive known, and he regarded Fabrosaurus as a nomen dubium potentially synonymous with Lesothosaurus due to overlapping material from the upper Elliot Formation.^[6] Thulborn (1992) countered by proposing Lesothosaurus as a junior synonym of the senior name Fabrosaurus, but this was largely rejected because the holotype of Fabrosaurus consists of insufficient material (a single jaw fragment), rendering it a nomen dubium, while Lesothosaurus is based on more diagnostic specimens.^[17] By the early 2000s, comprehensive reviews solidified Lesothosaurus's status as a key basal ornithischian outside Genasauria, with Weishampel and colleagues (in Norman et al., 2004) emphasizing its role in early ornithischian diversification while noting synonymy debates with other "fabrosaurs" like Technosaurus and Thescelosaurus, ultimately deeming Fabrosauridae obsolete as a formal taxon.^[17] Pre-2010 classifications often lumped it with contemporaneous South African basal ornithischians, including Heterodontosaurus, under the informal umbrella of "fabrosaurs," reflecting its importance in understanding Jurassic ornithischian radiation in Gondwana.^[17]

Phylogenetic position

Lesothosaurus diagnosticus is widely regarded as a basal ornithischian dinosaur, positioned outside the derived clade Genasauria, which unites cerapods and thyreophorans as more closely related to Triceratops horridus than to basal forms like Heterodontosaurus tucki. This placement reflects its retention of primitive ornithischian features, such as a simple dentition and unspecialized postcranial skeleton, distinguishing it from more advanced groups. Phylogenetic analyses consistently recover it near the base of Ornithischia, though its exact relationships remain debated, with some studies suggesting it as the sister taxon to all other ornithischians (excluding the even more basal Pisanosaurus mertii), while others nest it within Neornithischia as an early-diverging member. A key analysis incorporating postcranial data from multiple specimens positioned Lesothosaurus as a basal neornithischian, emphasizing characters like the elongate preacetabular process of the ilium and a fibula that is subequal in length to the tibia, which align it more closely with later neornithischians than with thyreophorans or cerapods. This 2016 study by Barrett, Butler, Knoll, and Porro highlighted how ontogenetic variation in the available material had previously obscured these traits, leading to unstable placements in earlier cladograms. However, the analysis underscored the fragility of such resolutions, as alternative character scorings could shift Lesothosaurus to a more basal position outside Neornithischia. Broader phylogenetic frameworks from the 2020s have occasionally supported thyreophoran affinities for Lesothosaurus, particularly in matrices emphasizing postcranial armor precursors or pelvic morphology, though these results are weakly supported and sensitive to taxon sampling. For instance, some datasets recover it as an early, unarmored thyreophoran due to shared primitive traits like a prominent acromial process on the scapula, which may represent a plesiomorphic condition rather than a synapomorphy. Debates persist regarding the monophyly of basal ornithischians (formerly "fabrosaurs"), with Lesothosaurus often serving as a wildcard taxon that challenges the stability of early divergences in Ornithischia. No close relatives have been confirmed, and the genus remains monospecific, with no significant phylogenetic revisions reported through 2025.

Paleobiology

Diet and feeding

Lesothosaurus diagnosticus exhibited a feeding strategy indicative of predominant herbivory supplemented by occasional omnivory, with adaptations for processing soft vegetation and possibly opportunistic consumption of small invertebrates or other matter.^[18]^[3] Its leaf-shaped, multicusped teeth, featuring marginal denticles, were suited for browsing on low-growing, non-abrasive plants such as shoots, fruits, and seeds, rather than grinding tougher foliage.^[18] A 2022 analysis of dental form and function confirmed a predominantly herbivorous diet for Lesothosaurus, likely composed of soft plant matter, with occasional omnivorous habits such as insect consumption.^[3] The predentary bone at the front of the lower jaw likely aided in precise nipping of vegetation, complementing the beak-like structure for selective foraging.^[19] Biomechanical modeling indicates Lesothosaurus possessed a relatively weak bite force, estimated at 121 N anteriorly and 258 N posteriorly, optimized for nipping and initial processing rather than forceful grinding or tearing.^[20] Given its adult body length of up to 2 meters, Lesothosaurus occupied an ecological niche that allowed exploitation of low-lying vegetation in the semi-arid paleoenvironment of the Early Jurassic upper Elliot Formation in southern Africa, where seasonal aridity limited plant availability to softer, seasonal growth.^[19]^[21] In comparison to more derived early heterodontosaurids like Heterodontosaurus, which developed stronger jaw muscles for higher bite forces and tougher plant matter, Lesothosaurus retained a more generalized feeding apparatus, reflecting its basal position among ornithischians and broader dietary flexibility.^[18]

Locomotion and behavior

Lesothosaurus was a facultatively bipedal dinosaur, with its reduced forelimbs—characterized by a forelimb-to-hindlimb length ratio of approximately 0.61—indicating they played a limited role in locomotion, likely serving for balance or minor manipulation rather than weight support.^[22] The hindlimbs were elongated and gracile relative to the body, featuring a near 1:1 ratio between femur and tibia lengths, adaptations that supported efficient cursorial movement suited to an agile lifestyle.^[23] These proportions, combined with a caudofemoralis-driven retraction mechanism, suggest Lesothosaurus could achieve moderate running speeds suitable for evasion.^[24]^[25] Taphonomic evidence from monodominant bonebeds containing multiple individuals across ontogenetic stages supports social aggregation and gregarious behavior in Lesothosaurus, with multigenerational herds inferred as a strategy for predator evasion in its Early Jurassic environment.^[26] Co-occurrence with larger carnivores such as Massospondylus in the Upper Elliot Formation implies that grouping facilitated escape from predation through collective vigilance and rapid flight.^[27] Bone histology reveals rapid vascularized growth during early ontogeny, slowing after 2–4 years, which likely promoted schooling among juveniles for enhanced protection during this vulnerable phase.^[26] However, no direct fossil evidence exists for nesting sites or parental care behaviors in Lesothosaurus.^[26]

Paleoenvironment

Geological setting

Lesothosaurus fossils are primarily known from the Upper Elliot Formation, which forms part of the Karoo Supergroup in the main Karoo Basin of southern Africa, encompassing outcrops in South Africa and Lesotho.^[28] This formation represents a continental red bed succession deposited during the Late Triassic to Early Jurassic transition.^[28] The lithology of the Upper Elliot Formation consists predominantly of red mudstones and fine- to medium-grained sandstones, indicative of a semi-arid floodplain environment with seasonal, ephemeral river systems and occasional aeolian influences.^[29] These sediments reflect low-energy depositional processes, including overbank flooding and channel lag accumulation, within a fluvio-lacustrine setting that transitioned from more perennial systems in the underlying Lower Elliot Formation.^[28] The regional climate during deposition was arid to semi-arid, characterized by highly seasonal rainfall and pedogenic alteration in the mudstones, suggesting periodic drought conditions.^[28] The age of the Upper Elliot Formation, and thus the Lesothosaurus-bearing horizons, is constrained to the Early Jurassic Hettangian–Sinemurian stages, approximately 201–190 Ma, based on U-Pb detrital zircon geochronology from tuffaceous sandstones and siltstones.^[28] Specific maximum depositional ages from analyzed samples range from <197.3 Ma to <191.9 Ma, confirming an Early Jurassic timeframe and distinguishing it from the underlying Norian–Rhaetian Lower Elliot Formation.^[28] Taphonomic evidence for Lesothosaurus indicates preservation in low-energy settings, such as channel lags and overbank deposits, where monodominant bonebeds formed through gradual accumulation rather than catastrophic events.^[30] These assemblages, often comprising multiple individuals across ontogenetic stages, suggest minimal transport and rapid burial in fine-grained sediments of the floodplain.^[30]

Associated fauna

The Upper Elliot Formation of South Africa and Lesotho, which has yielded the majority of Lesothosaurus fossils, preserves a moderately diverse Early Jurassic tetrapod assemblage dominated by archosaurs and synapsids, with limited evidence of other groups. Co-occurring dinosaurs include several basal sauropodomorphs, such as the abundant Massospondylus carinatus and rarer forms like Ignavusaurus rachelis and Ngwevu intloko, which were primarily herbivorous and may have competed with Lesothosaurus for low-lying vegetation.^[31]^[32] Heterodontosaurids, exemplified by Heterodontosaurus tucki, represent fellow basal ornithischians that likely occupied similar small-bodied herbivorous or omnivorous niches, potentially as direct competitors. Theropods are sparsely represented by body fossils but more commonly by tracks, including those of large carnivores (e.g., Moyenisauropus-like forms suggesting megatheropods over 8 m in length), indicating the presence of predators in the ecosystem. Overall, the formation has produced remains attributable to several dinosaur taxa, though many are fragmentary and the ornithischian component is particularly diverse compared to other Early Jurassic localities.^[33]^[34] Non-dinosaurian vertebrates further characterize this fauna as predator-rich and ecologically stratified. Non-mammaliaform cynodonts, notably the herbivorous tritylodontid Tritylodon cf. Rileyi, are among the most common elements, with over 100 specimens recovered, suggesting they formed a significant part of the small-to-medium herbivore guild alongside Lesothosaurus. Small crocodylomorphs, including Sphenosuchus acutus, Litargosuchus dundi, and Orthosuchus stormi, represent agile, terrestrial carnivores that likely preyed on smaller vertebrates, contributing to a dynamic food web. Rare occurrences of stereospondyl amphibians (e.g., brachyopoid jaw fragments), turtles (proganochelyid indet.), and lepidosaurs (sphenodontian-like) indicate semi-aquatic or riparian components, while pterosaur tracks (e.g., Pterosaurichnus-type) attest to aerial vertebrates in the community.^[35]^[36]^[37] Invertebrate remains are underrepresented in the upper Elliot Formation, reflecting its semi-arid depositional setting, but unionid bivalves (e.g., Unio sp.) occur in associated floodplain deposits, implying freshwater habitats nearby. Arthropod trace fossils, such as burrow casts, suggest the activity of insects or crustaceans in pedogenically altered sediments. Plant fossils are sparse due to the formation's aeolian-influenced paleoenvironments, but macroremains include horsetails (Equisetites sp.) and cycad-like fronds, while palynological data reveal a gymnosperm-dominated flora with abundant conifer (e.g., Podocarpus-type) and cycad pollen, alongside minor fern spores, supporting a low-diversity vegetation suited to seasonal aridity.^[38]^[39] Ecologically, Lesothosaurus occupied the niche of a small, agile herbivore or omnivore in a low-diversity assemblage where larger sauropodomorphs browsed higher vegetation and theropods/crocodylomorphs exerted predation pressure, fostering a community structured around floodplain resources amid a predator-rich landscape. A 2022 study on Lesothosaurus bonebeds further supports evidence of gregarious behavior in this fauna.^[30] No significant new faunal discoveries have emerged since the description of Ngwevu intloko in 2019, underscoring the stability of this paleoecological interpretation as of 2025.^[35]^[40]^[32]

References

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