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Plesiosaurus

Plesiosaurus is an extinct genus of sauropterygian marine reptile that lived during the Early Jurassic period, approximately 190 million years ago.^[1] The type and only valid species, P. dolichodeirus, is characterized by its long neck comprising about 40 cervical vertebrae, a small triangular skull with sharp conical teeth, a barrel-shaped body, four large paddle-like flippers adapted for underwater propulsion, and a relatively short tail.^[2] This species measured approximately 3 meters in length and inhabited shallow marine environments along what is now the coast of England.^[3] The genus was established based on fossils discovered in the late Sinemurian stage of the Lower Jurassic Charmouth Mudstone Formation near Lyme Regis, Dorset, England.^[4] The holotype specimen (NHMUK PV OR 22656), an articulated skeleton, was found by renowned fossil collector Mary Anning on December 10, 1823, and formally described by William D. Conybeare in 1824, who named it Plesiosaurus dolichodeirus meaning "near lizard with long neck."^[3] This discovery represented the first complete plesiosaur skeleton and provided key insights into the anatomy of these enigmatic reptiles, dispelling earlier misconceptions about their body plan.^[4] As a member of the family Plesiosauridae within Plesiosauroidea, Plesiosaurus exemplifies the "plesiosauromorph" body plan, with its elongated neck enabling ambush predation on schools of small fish, ammonites, and other invertebrates in coastal seas.^[5]^[2] Its flippers, derived from limbs with hyperphalangy (extra finger bones), allowed for efficient maneuvering akin to flying through water.^[5] Although over 25 partial to complete specimens of P. dolichodeirus are known, primarily from the UK, the genus highlights the diversity of early plesiosaurs and their role as apex predators in Mesozoic marine ecosystems before the group's radiation in later Jurassic and Cretaceous periods.^[1]

History of research

Discovery and initial description

The genus Plesiosaurus was established in 1821 by geologists William Daniel Conybeare and Henry Thomas De la Beche, based on a partial skeleton from the Lower Jurassic Lias Group in Dorset, England, originally in the collection of Colonel Thomas James Birch.^[6] In their preliminary paper presented to the Geological Society of London, they named the creature Plesiosaurus (from Greek plēsios, meaning "near," and sauros, "lizard"), interpreting it as a marine reptile intermediate in form between the recently described Ichthyosaurus and crocodilians, due to its elongated vertebral column and limb structures suggestive of aquatic adaptation.^[6] The first nearly complete articulated skeleton, which provided the basis for a more definitive understanding, was discovered by fossil collector Mary Anning on December 10, 1823, in the coastal cliffs near Lyme Regis, Dorset, England, also from the Lias Group.^[7]^[3] This specimen, measuring approximately 3 meters in length and remarkably well-preserved, was purchased by the Duke of Buckingham for 150 guineas and presented to the Geological Society of London, becoming the holotype for the type species Plesiosaurus dolichodeirus (meaning "long-necked").^[3] Conybeare provided the formal description in 1824, publishing "On the Discovery of an Almost Perfect Skeleton of the Plesiosaurus" in the Transactions of the Geological Society of London, where he detailed the animal's distinctive anatomy: a small head, a neck with around 35 vertebrae extending to nearly half the body's length, a compact trunk, a short tail, and four robust, paddle-like limbs for propulsion in marine environments. Accompanying sketches by De la Beche and Conybeare depicted Plesiosaurus as an agile swimmer with its neck arched upward, preying on fish in shallow seas, emphasizing its novelty compared to known reptiles.^[6] The creature's unprecedented morphology—particularly the disproportionately long neck—sparked initial confusion and debate among early 19th-century naturalists, who grappled with its affinities to Ichthyosaurus (sharing flipper-like limbs and marine habitat) while recognizing its lizard-like vertebral structure, ultimately affirming it as a new class of extinct aquatic reptile.^[6]

Subsequent studies and taxonomic revisions

In the decades following its initial description, numerous additional Jurassic fossils from localities across England, Germany, and other parts of Europe were referred to Plesiosaurus, often without rigorous comparison to the type material, resulting in the genus becoming a wastebasket taxon that encompassed a wide array of long-necked plesiosauroids.^[8] This over-referral included specimens from non-type localities, such as isolated vertebrae and partial skeletons attributed to new species like Plesiosaurus propinquus from the Toarcian of Yorkshire, which were later identified as representing distinct taxa due to discrepancies in vertebral counts and limb morphology.^[9] During the mid-20th century, studies began to address these issues through more detailed examinations of referred material. W.E. Swinton's work in the 1930s and 1940s, including descriptions of specimens like Macroplata tenuiceps (initially placed within Plesiosaurus), highlighted morphological variations in cervical vertebrae and helped refine identifications of early Jurassic forms, though many referrals persisted. Similarly, D.S. Brown's 1981 monograph on English Upper Jurassic plesiosauroids reviewed phylogenetic relationships and re-evaluated numerous specimens previously assigned to Plesiosaurus, synonymizing several species and restricting the genus to more diagnostic material based on shared autapomorphies such as the number of cervical vertebrae (approximately 40) and specific rib morphologies.^[10] Post-2000 revisions have further narrowed the scope of Plesiosaurus. Glenn W. Storrs's 1997 analysis clarified the genus's diagnostic features, emphasizing the type species P. dolichodeirus and excluding many historical referrals due to incomplete or mismatched anatomy, thereby reducing the number of valid species.^[8] More recently, Roger B.J. Benson and colleagues in 2012 conducted a taxonomic revision of the plesiosaurian assemblage from the Lower Jurassic of Street, Somerset, identifying multiple distinct taxa among fossils long attributed to Plesiosaurus and restricting the genus to specimens closely matching the holotype in size, neck length, and skull proportions, while erecting new genera for others.^[11] These efforts underscore ongoing recognition of historical errors, such as misattributions from distant localities lacking stratigraphic correlation to the type Sinemurian beds of Lyme Regis. While these revisions have stabilized the taxonomy, coverage remains somewhat outdated, with no major post-2020 publications directly addressing Plesiosaurus systematics; recent research has instead focused on new discoveries, such as basal plesiosauroids from the Posidonia Shale, suggesting potential for further updates as additional material is described.^[12]

Taxonomy

Classification within Plesiosauria

Plesiosaurus is classified within the clade Plesiosauria, a diverse group of extinct marine reptiles that arose during the Late Triassic and persisted until the end-Cretaceous mass extinction. Specifically, it occupies a basal position among plesiosauroids, the long-necked subgroup of Plesiosauria, and is assigned to the family Plesiosauridae, which encompasses early-diverging forms characterized by moderate neck lengths and generalized body plans. This placement reflects its role as one of the earliest well-known plesiosauroids, with the type species P. dolichodeirus serving as a foundational taxon for understanding the group's radiation.^[13] In evolutionary terms, Plesiosaurus originated in the Early Jurassic, approximately 200 million years ago, marking the initial diversification of plesiosauroids following the Late Triassic emergence of Plesiosauria. It shares close phylogenetic ties with sister taxa such as Microcleidus and other members of Plesiosauridae, forming a clade that diverged early from short-necked pliosauroids, including rhomaleosaurs, and from later long-necked lineages like elasmosaurs. This divergence highlights a key adaptive split within Plesiosauria, where plesiosauroids emphasized elongated necks for foraging in open waters, contrasting with the ambush predation strategies of pliosaurs. Phylogenetic analyses consistently recover Plesiosauridae as a monophyletic group at the base of Plesiosauroidea, though debates continue regarding the inclusion of certain transitional forms and the precise boundaries of the family.^[13]^[14] Seminal cladistic studies, notably Benson and Druckenmiller (2014), utilized extensive morphological datasets to position Plesiosaurus as a stem-plesiosauroid, supported by synapomorphies such as a moderate number of cervical vertebrae and specific humeral features that anchor its basal status. This analysis incorporated over 100 taxa and hundreds of characters, demonstrating robust support for Plesiosauridae's monophyly while revealing turnover events that shaped plesiosaur diversity across the Jurassic-Cretaceous boundary. More recent phylogenetic work, building on this framework, reaffirms Plesiosaurus's early-diverging role among plesiosauroids, with transitional taxa like Franconiasaurus bridging it to more derived cryptoclidians.^[13]^[14] Relative to outgroups within Sauropterygia, such as nothosaurs and pachypleurosaurs, Plesiosaurus exemplifies advanced marine adaptations that define Plesiosauria, including the modification of limbs into hydrofoil-like paddles and the expansion of limb girdles into propulsive platforms for efficient quadrupedal swimming in pelagic environments. These innovations distinguish plesiosaurs from their more amphibious sauropterygian ancestors, enabling fully aquatic lifestyles and global dispersal.

Valid species and nomenclature

The genus Plesiosaurus is monotypic, containing only its type species P. dolichodeirus Conybeare, 1824. This species is based on the holotype NHMUK PV OR 22656, an almost complete articulated skeleton discovered by Mary Anning at Lyme Regis, Dorset, England, in the Lower Jurassic Lias Group. The holotype, comprising a skull, most of the vertebral column (40 preserved cervical vertebrae), ribs, girdles, and limbs, was described as establishing the new genus and species based on its long neck, small head, and four equal-sized flippers. Numerous species originally assigned to Plesiosaurus have been reassigned to other genera following detailed morphological comparisons, reflecting ongoing taxonomic revisions. For example, P. brachypterygius Seeley, 1869, from the Lower Jurassic Posidonienschiefer Formation of Germany, was transferred to the new genus Hydrorion as H. brachypterygius based on unique features such as a short cervical vertebral column (26 vertebrae) and distinct humeral morphology differing from P. dolichodeirus. Similarly, P. guilelmiimperatoris Noetling, 1889, also from the Posidonienschiefer, was reclassified as Seeleyosaurus guilelmiimperatoris, reviving the genus Seeleyosaurus Dames, 1897, due to differences in axial skeleton proportions and cranial elements. Other former species include P. cramptoni Tate and Blake, 1876, now regarded as a junior synonym of Rhomaleosaurus cramptoni (Seeley, 1874), a pliosauroid distinguished by its shorter neck and larger skull. The species P. macrocephalus Stutchbury, 1846, remains under review but is excluded from Plesiosaurus pending further analysis of its juvenile holotype and potential affinities with basal plesiosauroids like Anningasaura.^[15]^[16] Nomenclatural stability for Plesiosaurus has been reinforced through recent systematic revisions, including those by Druckenmiller and Benson (2012), which affirm P. dolichodeirus as the sole valid species while reassigning others to avoid taxonomic wastebaskets. No specific ICZN rulings have been required for the genus name, as it adheres to priority and description standards established in the original publication. Species delimitation within Plesiosauria relies on comparative morphology (e.g., vertebral counts, limb proportions, and cranial sutures), ontogenetic stage (to distinguish juveniles from adults), and stratigraphic locality, ensuring referrals match the Early Jurassic European context of the type material. These criteria highlight P. dolichodeirus as a distinct basal plesiosauroid with 40 cervical vertebrae and a body length of approximately 3–4 meters.^[11]

Anatomy

Skull and dentition

The skull of Plesiosaurus dolichodeirus is small and triangular in outline, measuring approximately 18–25 cm in length, with a narrow profile that achieves maximum width at the postorbital bar. The anterior portion forms a bluntly triangular rostrum, while the temporal region is elongated, accommodating large, quadrangular temporal fenestrae that house the jaw adductor muscles. The orbits are prominent, roughly circular, and positioned near the midpoint of the skull length, facing slightly dorsolaterally to enhance binocular vision in a marine environment. External nares are small and positioned anterior to the orbits, and the pineal foramen is relatively large, positioned between the orbits on the midline. These features are evident in the type specimen NHMUK PV OR 22656, a nearly complete skeleton from the Lower Jurassic Lias of Lyme Regis, England, where the skull measures about 18.5 cm long.^[17]^[18]^[19] Dentition in P. dolichodeirus consists of homodont, conical teeth that are sharp-pointed, slightly recurved, and circular to oval in cross-section, adapted for grasping soft-bodied prey. Each maxillary tooth row bears approximately 18–20 teeth, with the premaxilla contributing 4–5 smaller teeth anteriorly; the lower jaw has a comparable number, including up to 4 teeth in the symphyseal region. Tooth crowns are slender, up to 2–3 cm long, with fine longitudinal ridges on the enamel surface for added grip, and roots that fit into deep alveolar sockets. Jaw mechanics involve a simple gape-and-grasp mechanism, with the elongated temporal region supporting moderate adductor leverage, though specific bite force estimates remain unquantified due to limited cranial material. In the type specimen NHMUK PV OR 22656, the dentition aligns with this pattern, showing uniform conical morphology without significant heterodonty.^[17]^[20] Sensory adaptations in the skull include the large orbits, suggesting reliance on acute vision for detecting prey in open water, and the prominent pineal foramen, which likely housed a parietal eye for light detection and circadian regulation. The nares, though externally small, connect to internal choanae positioned posteriorly, potentially aiding in chemosensory olfaction through water flow across olfactory epithelia. These cranial features reflect adaptations to a fully aquatic lifestyle, distinct from terrestrial reptiles, as observed across multiple P. dolichodeirus specimens including the type NHMUK PV OR 22656.^[17]^[21]^[22]

Vertebral column and axial skeleton

The vertebral column of Plesiosaurus dolichodeirus comprises approximately 38–42 cervical vertebrae, 21 dorsal vertebrae, 2–3 sacral vertebrae, and 28 caudal vertebrae, yielding a total of about 90 vertebrae and emphasizing the disproportionately long neck relative to the short tail.^[1] The cervical series transitions gradually into the dorsal series via 3–5 pectoral vertebrae, where rib facets shift from the centra to elongate transverse processes on the neural arches.^[23] Overall, the axial skeleton accounts for roughly half of the animal's total body length of 3–3.5 m.^[1] Cervical vertebrae are amphicoelous to platycoelous, with relatively elongate centra measuring 4–5 cm in length, collectively forming a neck up to 2 m long that provides structural support and flexibility through overlapping zygapophyses.^[23]^[24] These vertebrae feature paired subcentral foramina on the ventral surfaces, while the neural arches bear small, rugose diapophyses for rib articulation.^[25] Dorsal vertebrae are shorter and more robust, with high neural spines and broad capitular and tubercular facets for single-headed ribs that enhance body rigidity and contribute to buoyancy regulation.^[23] Sacral vertebrae are fused to the pelvis, and caudal vertebrae taper rapidly, supporting a abbreviated tail without chevrons in the proximal region.^[1] Cervical ribs are short, hatchet-shaped, and double-headed, articulating via distinct capitulum and tuberculum facets to allow neck mobility without restricting movement.^[23] Dorsal ribs are long, thick, and single-headed, flaring distally to form a robust thoracic cage, while gastralia consist of tightly interlocked, boomerang-shaped elements arranged in rows between the pectoral and pelvic girdles, providing ventral body support.^[1]^[23] This configuration of the axial skeleton underscores P. dolichodeirus's adaptation for a streamlined body form, with the flexible neck playing a minor role in propulsion compared to the limbs.^[26]

Limbs and girdles

The pectoral girdle of Plesiosaurus dolichodeirus consists of broad scapulae and large coracoids forming expanded, flat ventral plates that meet medially in a symphysis, providing extensive surfaces for muscle attachments to support the forelimbs.^[23] The scapulae are diamond-shaped with a reduced dorsal blade, while the coracoids are broad plates contributing significantly to the anterolaterally oriented glenoid fossa for humeral articulation.^[1] In the pelvic girdle, the ilia are small, rod-like elements measuring approximately 10.7–10.9 cm in length, expanded at the extremities with a twisted shaft connecting to the sacral ribs and ischium; the pubis and ischium form large, flat ventral plates with a medial symphysis for hindlimb support.^[23]^[1] The limbs are modified into four elongated, paddle-like flippers adapted for aquatic locomotion, with the forelimbs longer and more robust than the hindlimbs, reflecting asymmetry where forelimbs serve as primary propulsors and hindlimbs as auxiliaries.^[27] In a well-preserved specimen, the humerus reaches 21.4–21.5 cm in length, exceeding the femur at 18.4–18.6 cm, while the full forepaddle extends to approximately 70 cm and the hindpaddle to 50 cm based on reconstructed proportions from multiple fossils.^[1] The humerus is robust with a rounded proximal head, oval midshaft, and expanded distal epicondyles; epipodials (radius and ulna) are shortened (7.9–8 cm) and broadened to increase paddle surface area, often covered by cartilaginous caps in life.^[28] Hyperphalangy is evident in the autopodia, with at least 9 phalanges preserved in hindlimb digits of some specimens, though complete counts suggest 10–12 per digit in the outer rays for enhanced flipper flexibility.^[1] These structural features contribute to buoyancy control during swimming.^[29]

Paleobiology

Locomotion and buoyancy

Plesiosaurus, like other plesiosauroids, propelled itself through water using a form of underwater flight powered primarily by its enlarged forelimbs, which functioned as hydrofoils to generate lift and thrust through dorso-ventral oscillations.^[30] The hindlimbs, smaller than the forelimbs, played a secondary role in steering and fine maneuvering, contributing additional thrust when synchronized with the forelimbs to enhance overall efficiency by up to 40%.^[31] The long neck likely served as a stabilizer or rudimentary rudder, aiding in balance during turns without requiring extreme lateral flexion.^[32] Buoyancy in Plesiosaurus was maintained through a combination of dense bones that increased overall body density to approach neutral buoyancy at depth, complemented by air-filled lungs that provided positive buoyancy when surfaced.^[33] Computational models indicate that with full lungs, the animal would float at the surface, but as air was compressed during dives, the dense skeletal structure prevented excessive sinking, allowing sustained submergence without constant swimming effort.^[34] Surfacing behaviors, inferred from lung capacity and bone density, suggest periodic breaths to replenish oxygen stores, enabling dives of moderate duration.^[32] Cruising speeds for Plesiosaurus have been estimated at 0.5–1 m/s based on hydrodynamic simulations incorporating limb proportions, body mass of approximately 200–300 kg, and drag coefficients derived from its streamlined form.^[30] These velocities align with energy-efficient locomotion for a small-bodied plesiosauroid, sufficient for patrolling shallow marine environments without rapid fatigue.^[35] Debates persist regarding neck flexibility in Plesiosaurus, with anatomical analyses suggesting a relatively stiff posture rather than the highly maneuverable S-shape once proposed; instead, a static, streamlined alignment minimized drag during forward swimming.^[32] This interpretation, supported by vertebral zygapophyseal facets and muscle attachment sites, implies the neck's primary role was hydrodynamic stability over agile predation strikes, though the issue remains unresolved pending further fossil evidence.^[36]

Diet and feeding ecology

Plesiosaurus was primarily a piscivorous predator, with its diet inferred from tooth morphology and rare direct fossil evidence. The genus possessed small, pointed teeth with longitudinal striations, characteristic of soft-prey generalists adapted for grasping fish and soft-bodied invertebrates like cephalopods, rather than crushing hard-shelled prey. Direct evidence of diet comes from a juvenile Plesiosaurus sp. specimen from the Lower Jurassic (Pliensbachian) Lias Group near Charmouth, Dorset, England, where disarticulated fish skeletal remains were preserved within the abdominal region, interpreted as undigested stomach contents indicating a recent meal of small fish.^[37] Coprolites from Early Jurassic marine deposits, likely produced by plesiosaurs based on size, shape, and associated fauna, contain fish scales, belemnite guards, cephalopod hooklets, and fragments of bivalves and gastropods, supporting a diet that included fish, cephalopods, and occasional benthic mollusks. The hunting strategy of Plesiosaurus is reconstructed as that of an ambush predator, leveraging its elongated neck to extend the head rapidly toward prey while the body remained relatively stationary to minimize detection in the water column. This approach, combined with the small skull size and needle-like dentition, was well-suited for capturing elusive, soft-bodied prey such as schooling fish or swimming cephalopods without the need for sustained pursuit. Within Jurassic marine food webs, Plesiosaurus functioned as a mid-level carnivore, occupying a trophic position above primary consumers like fish and cephalopods but below apex predators, and likely competed for similar resources with contemporaneous ichthyosaurs that exhibited overlapping piscivorous habits.^[38]

Reproduction and growth

The reproductive biology of Plesiosaurus is inferred primarily from evidence in related plesiosaurs, as no direct fossils of embryos or neonates attributable to this genus have been discovered. Viviparity, or live birth, is supported by a Late Cretaceous specimen of the plesiosaur Polycotylus latipinnis preserving a near-term fetus within a gravid female, indicating internal gestation and birth of a single large offspring in water, consistent with the group's fully aquatic adaptations and inability to haul out on land.^[39] This K-selected strategy, featuring low fecundity but high parental investment, is extrapolated to Plesiosaurus and other Jurassic plesiosauroids, with no fossil eggs known from the entire Plesiosauria clade to suggest oviparity.^[29] Growth patterns in Plesiosaurus reflect rapid early ontogeny typical of plesiosaurs, as revealed by bone histology. Femoral sections exhibit fibrolamellar bone tissue with radial primary osteons, signaling elevated metabolic rates and accelerated somatic growth comparable to those of extant endothermic vertebrates.^[40] Quantitative analyses of related taxa indicate juveniles achieved substantial size—up to 60% of adult body length—within the first few years postnatally, with overall maturation involving few annual growth cycles before an external fundamental system formed to halt appositional growth.^[29] For Plesiosaurus, reaching an adult length of approximately 3–5 meters, this implies a lifespan potentially spanning decades, though precise age-at-maturity estimates remain elusive without scleral ring data specific to the genus. Sexual dimorphism in Plesiosaurus is poorly documented and unconfirmed, with no consistent morphological differences (such as in paddle proportions) identified in available specimens that cannot be attributed to ontogenetic variation or preservation artifacts. Stratigraphic distributions of plesiosaur fossils in nearshore Jurassic deposits suggest possible breeding migrations to shallower marine environments for parturition, but this remains speculative without direct ontogenetic series. Current understanding is limited by the scarcity of Plesiosaurus juvenile material and reliance on congeners like Polycotylus, highlighting significant gaps in direct evidence for reproductive behaviors and life history in this basal plesiosauroid.^[29]

Paleoenvironment and distribution

Geological formations

Fossils of Plesiosaurus are primarily known from the Sinemurian Stage of the Lower Jurassic, occurring in the Blue Lias Formation and the overlying Charmouth Mudstone Formation, both part of the Lias Group in Dorset, England, dating to approximately 199.5–192.9 million years ago.^[41] These formations represent the basal units of the Lias Group, with the Blue Lias Formation spanning the latest Hettangian to early Sinemurian and the Charmouth Mudstone extending into the Pliensbachian.^[42] The type species P. dolichodeirus was first described from specimens collected at Lyme Regis in these strata.^[43] The lithology of these formations consists predominantly of interbedded mudstones, shales, and limestones, deposited in a shallow epicontinental sea environment characterized by low-energy marine conditions.^[44] The Blue Lias features rhythmic alternations of thin limestones and calcareous mudstones, while the Charmouth Mudstone is more mudstone-dominated with occasional limestone nodules, reflecting periodic changes in sedimentation influenced by sea-level fluctuations and climate.^[45] These deposits formed in a subtropical shelf sea, with lagoonal and restricted basin settings promoting fine-grained sedimentation.^[46] Taphonomic conditions in these formations favored exceptional preservation of Plesiosaurus skeletons, often found fully articulated due to anoxic bottom waters that inhibited scavenging and bioturbation.^[45] Fossils are typically embedded in pyritized or calcareous concretions within the mudstones, preserving delicate structures like vertebrae and paddle elements with minimal distortion; disarticulation is rare, as evidenced by complete specimens from lagoonal facies. Key sites include the coastal exposures at Lyme Regis, where wave action erodes the cliffs to reveal fossils, and inland quarries at Street in Somerset, also within the Blue Lias Formation.^[47] These strata encompass the Hettangian to Pliensbachian interval of the Lower Jurassic, providing a stratigraphic framework for Plesiosaurus occurrences.^[48]

Geographic and stratigraphic range

Plesiosaurus is restricted to the Early Jurassic Epoch, with a stratigraphic range spanning the Sinemurian and Pliensbachian stages, encompassing a duration of approximately 5 to 10 million years.^[4]^[49] The genus is best documented from the late Sinemurian substage in the Charmouth Mudstone Formation of the Lower Lias Group, though fragmentary remains suggest extension into the lower Pliensbachian in some localities.^[50] This temporal distribution aligns with the initial radiation of plesiosauroids following their appearance in the latest Triassic, during a period of expanding shallow marine environments across the Tethyan realm.^[51] Geographically, Plesiosaurus exhibits a limited distribution centered on the southern coast of England, particularly in Dorset (e.g., Lyme Regis and Charmouth) and Somerset (e.g., Street), where the majority of articulated skeletons have been recovered from coastal exposures of the Lower Lias.^[52]^[53] Possible referrals extend the range to nearby European sites, including isolated elements from northern France (Normandy) and Germany (North Rhine-Westphalia and Lower Saxony), though these assignments remain tentative pending further taxonomic revision.^[54] Overall, the genus appears endemic to the subtropical margins of the Tethys Ocean, reflecting a localized adaptation to epicontinental seaways during the early Mesozoic marine transgression.^[55] In terms of paleoecology, Plesiosaurus inhabited warm, shallow shelf seas characterized by soft mud substrates and moderate oxygenation, where it co-occurred with diverse marine invertebrates such as ammonites (e.g., Arieticeras) and bivalves (e.g., Gryphaea), as well as other reptiles including ichthyosaurs like Ichthyosaurus communis. These associations indicate a shared habitat in productive coastal ecosystems supportive of a fish-rich food web. Despite its presence in productive lagerstätten, Plesiosaurus was relatively uncommon, with around 20 known specimens—mostly partial skeletons—suggesting lower abundance compared to contemporaneous plesiosauroids or ichthyosaurs in the same formations.^[4]^[56]

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All cervical vertebrae of plesiosaurs show a pair of large foramina on the ventral surface of the vertebral centrum, called subcentral foramina or subcentralia ...
[27]
Neck function - Plesiosaur Directory
The holotype of Plesiosaurus is a typical example of this straight-necked posture. Holotype specimen (NHMUK PV OR 22656) of Plesiosaurus dolichodeirus on ...Missing: skull measurements
[28]
Ecomorphology of plesiosaur flipper geometry - ResearchGate
Aug 10, 2025 · Plesiopterys possesses a high count of cervical vertebrae and a relatively small head, yet the limbs and limb girdles are plesiomorphic in ...
[29]
Determination of muscle strength and function in plesiosaur limbs
Jun 3, 2022 · The Plesiosauria (Sauropterygia) are secondary marine diapsids. They are the only tetrapods to have evolved hydrofoil fore- and hindflippers ...
[30]
Plesiosaurs - ScienceDirect.com
May 22, 2023 · Large marine reptile resembling a large sea turtle, but with four evenly shaped flippers and looking as though a large snake had been pulled through its ...
[31]
Computer Simulations Imply Forelimb-Dominated Underwater Flight ...
Neutral limb positions were derived from existing estimates for Plesiosaurus sp. [3]. In the forelimb the neutral position is -15 degrees from the ...
[32]
The four-flipper swimming method of plesiosaurs enabled efficient ...
Aug 30, 2017 · We show that plesiosaur hind flippers generated up to 60% more thrust and 40% higher efficiency when operating in harmony with their forward counterparts.Abstract · Introduction · Material and methods · Results and discussion
[33]
[PDF] An integrated approach to understanding the role of the long neck in ...
Mar 1, 2017 · Plesiosaurus dolichodeirus swimming at the water surface, possibly ... number of cervical vertebrae anteriorly to prevent the neck from ...
[34]
Floating point: a computational study of buoyancy, equilibrium, and ...
Mar 12, 2007 · The idea that plesiosaurs could maintain their necks above the water surface in an erect manner is rejected due to unbalanced buoyancy torques ...
[35]
A computational study of buoyancy, equilibrium, and gastroliths in ...
It has been argued, from analysis of mathematical models of floating plesiosaurs (Henderson 2006) , that gastroliths had little impact on buoyancy. However, a ...
[36]
(PDF) Plesiosaur Swimming as Interpreted from Skeletal Analysis ...
Aug 5, 2025 · The most efficient underwater swimming was synchronous and semi-synchronous movements of the fore and hind flippers.
[37]
An integrated approach to understanding the role of the long neck in ...
Aug 5, 2025 · ... neck in plesiosaurs, and how the problems associated with stiffness or flexibility ... Noè et al., 2017; Wintrich et al., 2017b). Storrs (1991) In ...
[38]
(PDF) Bottom-Feeding Plesiosaurs - ResearchGate
Aug 6, 2025 · Some mosasaurs have their stomach contents preserved, providing direct and unambiguous evidence of diet. The results show a relatively clear ...
[39]
Macropredatory ichthyosaur from the Middle Triassic and the origin ...
Jan 7, 2013 · Here we report a top-tier tetrapod predator, a very large (>8.6 m) ichthyosaur from the early Middle Triassic (244 Ma), of Nevada.Missing: carnivore | Show results with:carnivore
[40]
Atmospheric Blocking and Atlantic Multidecadal Ocean Variability
**Summary:**
[41]
Evolutionary implications of the divergent long bone histologies of ...
Jun 18, 2013 · The Plesiosaurus femur appears to be osteoporotic in some regions of the cortex but not in all. Osteoporosis is not as prevalent as shown by ...Tetrapod Long Bone Histology... · Methods · Discussion
[42]
A marine reptile fauna from the Early Jurassic Saltford Shale (Blue ...
Aug 7, 2025 · ... Blue Lias and Charmouth Mudstone formations (Page 2010) . ... Plesiosaurus dolichodeirus and its junior synonyms can be assigned to Plesiosaurus.
[43]
The Early Jurassic sequence of Lyme Regis, Dorset, England and its ...
Dec 20, 2023 · Lyme Regis exposes Early Jurassic marine limestones and mudstones, including the Blue Lias and Charmouth Mudstone Formations, and is known for ...
[44]
'THE MOST WONDERFUL WONDER EVER DISCOVERED': MARY ...
... Blue Lias of Lyme Regis. Anning's 1823 Plesiosaurus discovery established her fame and reputation as a fossilist. ResearchGate Logo. Discover the world's ...
[45]
Lias Group - British Geological Survey
The Lias Group consists mainly of clays, mudstones and limestones of Late Triassic and Early Jurassic age, deposited between 180 and 205 million years ago.Missing: Charmouth | Show results with:Charmouth
[46]
Combined sea-level and climate controls on limestone formation ...
Jan 30, 2017 · The Blue Lias Formation is well known for fully articulated ichthyosaur and plesiosaur fossils (e.g. Milner & Walsh, 2010). Skeletons enclosed ...
[47]
The Early Jurassic sequence of Lyme Regis, Dorset, England and its ...
The cliffs and foreshore at Lyme Regis on the Dorset coast expose a sequence of Early Jurassic marine limestones and mudstones of the Blue Lias Formation ...
[48]
Fossil Collection - Alfred Gillett Trust
The collection includes marine reptiles, such as Ichthyosaurs and a plesiosaur, from the Blue Lias quarries of Street, Somerset, and is now under the Alfred ...
[49]
A Plesiosaur Specimen from the Lias of Lyme Regis - ResearchGate
Aug 5, 2025 · ... Blue Lias Formation and the overlying Charmouth Mudstone Formation, the lowest units of the Hettangian-Aalenian Lias Group. Known for its ...
[50]
(PDF) A New Pliosaurid from the Pliensbachian, Early Jurassic of ...
Aug 5, 2025 · This paper offers a re-description of one of the oldest (Lower Hettangian, Lower Jurassic) plesiosaurians (Reptilia: Sauropterygia) from Germany ...
[51]
[PDF] New plesiosaur fossils from the Pliensbachian (Early Jurassic) of ...
Aug 4, 2025 · New plesiosaur fossils were found in Bornholm, including the first juvenile specimen from Denmark, and the first of its kind found in Denmark.
[52]
[PDF] A new plesiosaur from the Lower Jurassic of Portugal and the early ...
Jun 9, 2021 · Carbon isotope stratigraphy and ammonite biochronostratig- raphy across the Sinemurian–Pliensbachian boundary in the western. Iberian margin.
[53]
[PDF] A Juvenile Specimen of ?Plesiosaurus sp. from the Lias (Lower ...
Twenty-three vertebrae, and a small portion of a twenty-fourth, are preserved in the body nodules for a total known vertebral count of forty-four. Distinction ...
[54]
On a New Species of Plesiosaurus from Street, near Glastonbury
The specimen is a remarkably fine one. The limestone matrix in which it is imbedded being hard and free from pyrites, every part is well preserved ; and the ...Missing: formation | Show results with:formation
[55]
A New Pliosaurid from the Pliensbachian, Early Jurassic of ...
Feb 7, 2012 · Complete plesiosaur skeletons are relatively abundant in the Lower Jurassic of England and Germany, but are exceptional in France.
[56]
Early-diverging plesiosaurs from the Pliensbachian (Lower Jurassic ...
Nov 26, 2024 · Here, we describe two previously unreported lower Pliensbachian plesiosaur occurrences that originate from two sites located in North Rhine-Westphalia, Germany.Missing: range | Show results with:range
[57]
(PDF) A Juvenile Plesiosaur from the Pliensbachian (Lower Jurassic ...
Aug 10, 2025 · Pending histological analysis, plesiosaur specimens previously considered to be juveniles may simply be paedomorphic adults, unless ...