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Pliosauroidea
Pliosauroidea
Pliosauroidea
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Pliosauroids
Temporal range: Hettangian–Turonian
Cast of Rhomaleosaurus cramptoni (NHMUK PV R 34), Natural History Museum, London
Liopleurodon ferox mounted skeleton, Museum of Paleontology, Tübingen
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Superorder: †Sauropterygia
Order: †Plesiosauria
Clade: †Neoplesiosauria
Suborder: †Pliosauroidea
Welles, 1943
Families and genera

see text

Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive toothed jaws, commonly known as pliosaurs. More primitive non-thalassophonean pliosauroids resembled plesiosaurs in possessing relatively long necks and smaller heads. They originally included only members of the family Pliosauridae, of the order Plesiosauria, but several other genera and families are now also included, the number and details of which vary according to the classification used.

The distinguishing characteristics are a short neck and an elongated head, with larger hind flippers compared to the fore flippers, the opposite of the plesiosaurs. They were carnivorous and their long and powerful jaws carried many sharp, conical teeth. Pliosaurs range from 4 to 10 meters or more in length.[1][2] Their prey may have included fish, sharks, ichthyosaurs, dinosaurs and other plesiosaurs.

The largest known species are Kronosaurus and Pliosaurus macromerus; other well known genera include Rhomaleosaurus, Peloneustes, and Macroplata.[3] Fossil specimens have been found in Africa, Australia, China, Europe, North America and South America.

Many very early (from the Early Jurassic and possibly Latest Triassic, i.e. Rhaetian) primitive pliosauroids were very like plesiosauroids in appearance and, indeed, used to be included in the family Plesiosauridae.

Name

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Liopleurodon ferox

Pliosauroidea was named by Welles in 1943. It is adapted from the name of the genus Pliosaurus, which is derived from the Greek πλειων (pleion), meaning "more/closely", and σαυρος (sauros) meaning "lizard"; it therefore means "more saurian". The name Pliosaurus was coined in 1841 by Richard Owen, who believed that it represented a link between plesiosauroids and crocodilians (considered a type of "saurian"), particularly due to their crocodile-like teeth.


Classification

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Taxonomy

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Macroplata
Pliosaurus
Simolestes vorax
Attenborosaurus

The taxonomy presented here is mainly based on the plesiosaur cladistic analysis proposed by Hilary F. Ketchum and Roger B. J. Benson, 2011 unless otherwise noted.[4]

Phylogeny

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Cast of "Plesiosaurus" macrocephalus found by Mary Anning, Muséum national d'histoire naturelle, Paris

Pliosauroidea is a stem-based taxon that was defined by Welles as "all taxa more closely related to Pliosaurus brachydeirus than to Plesiosaurus dolichodeirus". Pliosauridae and Rhomaleosauridae are stem-based taxa too. Pliosauridae is defined as "all taxa more closely related to Pliosaurus brachydeirus than to Leptocleidus superstes, Polycotylus latipinnis or Meyerasaurus victor". Rhomaleosauridae is defined as "all taxa more closely related to Meyerasaurus victor than to Leptocleidus superstes, Pliosaurus brachydeirus or Polycotylus latipinnis".[8] The cladogram below follows a 2011 analysis by paleontologists Hilary F. Ketchum and Roger B. J. Benson, and reduced to genera only.[4]

Pliosauroidea

Large pliosauroids

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In 2002, the discovery of a very large pliosauroid was announced in Mexico. This pliosauroid came to be known as the "Monster of Aramberri". Although widely reported as such, it does not belong to the genus Liopleurodon.[10] The remains of this animal, consisting of a partial vertebral column, were dated to the Kimmeridgian of the La Caja Formation.[11] The fossils were found much earlier, in 1985, by a geology student and were at first erroneously attributed to a theropod dinosaur by Hahnel.[12] The remains originally contained part of a rostrum with teeth (now lost).

In August 2006, palaeontologists of the University of Oslo discovered the first remains of a pliosaur on Norwegian soil. The remains were described as "very well preserved, as well as being unique in their completeness". The large animal was determined to be a new species of Pliosaurus.[3] In the summer of 2008, the fossil remains of the huge pliosaur were dug up from the permafrost on Svalbard, a Norwegian island close to the North Pole.[13] The excavation of the find is documented in the 2009 History television special Predator X.

On 26 October 2009, palaeontologists reported the discovery of potentially the largest pliosauroid yet found. Found in cliffs near Weymouth, Dorset, on Britain's Jurassic Coast, the fossil had a skull length of 6 feet 5 inches (1.96 meters). Palaeontologist Richard Forrest told the BBC: "I had heard rumours that something big was turning up. But seeing this thing in the flesh, so to speak, is just jaw dropping. It is simply enormous."[14] It was determined that the specimen belonged to a new species that scientists named Pliosaurus kevani.[15]

In December 2023, the recent discovery of a pliosaur skull on the Dorset coast was described as "one of the most complete specimens of its type ever discovered".[16] The discovery and research of the skull was covered in the PBS documentary Attenborough and the Jurassic Sea Monster hosted by David Attenborough.[17]

References

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Pliosauroidea

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Pliosauroidea is an extinct of plesiosaurian marine reptiles within the larger group , characterized by their short necks, large skulls, and robust bodies adapted for powerful predation in oceans. These reptiles, often referred to as pliosaurs, featured a "pliosauromorph" with elongated snouts, conical or trihedral teeth for gripping prey, and strong muscles capable of generating immense bite forces, distinguishing them from the long-necked plesiosauroids. Their fossils are known from deposits spanning the to the early , a temporal range of approximately 120 million years during which they occupied diverse marine environments worldwide. Phylogenetically, Pliosauroidea forms a monophyletic group sister to within Neoplesiosauria, comprising basal taxa and derived families such as Rhomaleosauridae (early forms with relatively longer necks) and (advanced, giant predators). The clade's diversity peaked in the Middle to , with the pliosauromorph body plan evolving independently at least three times, reflecting adaptive convergence rather than a single linear progression. Notable genera include , a giant with skulls up to 2 meters long and estimated body lengths exceeding 10 meters, exemplifying the group's role as apex predators that preyed on fish, ammonites, and other marine reptiles. Pliosaurids like those in exhibited high bite forces ranging from 9,600 to 48,000 Newtons, enabling them to crush bone and dismember large prey in ecosystems such as the Formation. By the , Pliosauroidea diversified further, with forms like those in the Pacific extending into the stage, before declining toward the end of the , leading to their extinction by the early . Their record, preserved in lagerstätten across , , and , provides critical insights into the evolutionary dynamics of marine tetrapods and the structure of ancient food webs.

Etymology and Overview

Name

Pliosauroidea is a clade of extinct marine reptiles within Plesiosauria, named as a superfamily by paleontologist Samuel P. Welles in 1943. The term derives from the genus Pliosaurus, which was established by Richard Owen in 1841 to describe Jurassic plesiosaur specimens from England, particularly those exhibiting robust cranial features. Etymologically, Pliosaurus combines the Greek words pleion (πλεῖον, meaning "more") and sauros (σαῦρος, meaning "lizard"), originally intended to signify "more saurian" in reference to the genus's perceived resemblance to crocodilian reptiles rather than the slender-necked Plesiosaurus. Historically, the nomenclature evolved from Owen's initial description of Pliosaurus as a distinct genus within Plesiosauria, separating it from long-necked forms based on morphological differences observed in British fossils. Early classifications treated Pliosauroidea as a family-level group (Pliosauridae), but by the mid-20th century, Welles formalized it as a superfamily encompassing short-necked plesiosaurs. In modern taxonomy, it has transitioned to a clade-based framework, reflecting phylogenetic analyses that emphasize evolutionary relationships over traditional Linnaean ranks. Welles provided a stem-based definition for Pliosauroidea: all plesiosaurs more closely related to Pliosaurus brachydeirus than to Plesiosaurus dolichodeirus. This definition has been retained in subsequent revisions, anchoring the clade to its while accommodating diverse morphologies across marine ecosystems.

General Description

Pliosauroidea is an extinct of marine reptiles within Plesiosauria, distinguished by their short necks comprising 10 to 20 , in contrast to the longer necks (often exceeding 30 vertebrae) of the sister . These short-necked plesiosaurs featured large, robust skulls up to 2 meters in length, powerful jaws lined with conical teeth, elongated bodies, and four paddle-like flippers adapted for underwater propulsion. Overall body lengths ranged from approximately 4 meters in smaller species to over 13 meters in the largest forms, with the heaviest individuals estimated to exceed 20 metric tons in mass. Fossils of pliosauroids document a temporal range from the Hettangian stage to the early Turonian stage, encompassing roughly 130 million years of marine history. Their remains have been recovered from deposits across all continents except , reflecting a in ancient oceans worldwide. As carnivorous apex predators, pliosauroids dominated marine food webs, utilizing their massive and bite forces to capture and subdue prey including , cephalopods, and other marine reptiles. Many advanced members belonged to the subclade Thalassophonea, which emphasized hypercarnivorous adaptations such as enlarged skulls and enhanced feeding mechanics for tackling large vertebrate prey.

Anatomy

Cranial Features

The skulls of pliosauroids exhibit specialized morphology adapted for macropredation, with variations reflecting evolutionary trends across the clade. Early pliosauroids, such as those in Rhomaleosauridae, typically feature an elongated rostrum that constitutes a significant proportion of skull length, often exceeding 50% of the total cranial dimension, facilitating prey detection and capture in diverse marine environments. In contrast, later pliosaurids display a shortened, more robust rostrum relative to overall size, with preorbital regions comprising around 57% of length in forms like Pliosaurus kevani, emphasizing bite force over reach. Large temporal fenestrae are ubiquitous, providing extensive attachment surfaces for adductor musculature and enabling powerful closing forces estimated at 27,685 to 48,728 Newtons in advanced pliosaurids like P. kevani. Orbits are positioned dorsally and anteriorly, promoting some degree of to aid in precise targeting of prey during hunts. Dental adaptations in pliosauroids underscore their role as apex predators, with conical, interlocking teeth designed for piercing and holding prey. These teeth feature subtrihedral or trihedral cross-sections in derived forms, with sparse to coarse apicobasal ridges on the enamel for enhanced grip, and crowns that curve lingually to prevent escape of struggling victims. In gigantic pliosaurids like Pliosaurus funkei, the teeth are large, reflecting adaptations for tackling prey. Tooth replacement occurs continuously, with resorption pits and developing successors visible in histological sections, allowing sustained functionality despite high wear; exhibit longer replacement cycles and symmetrical patterns, while posterior ones show shorter, asymmetrical cycles. Wear patterns on crowns, including flat facets and striations, indicate capability for bone-crushing in robust pliosaurid taxa, distinguishing them from more piercing-focused dentitions in earlier relatives. Although primarily a single marginal row, the interlocking nature and replacement dynamics create functional multiplicity, as seen in Liopleurodon where dense alveolar packing supports prolonged engagement with prey. Sensory adaptations in pliosauroid crania enhance underwater hunting efficiency. Large external nares, positioned anteriorly near the rostrum tip, suggest improved olfactory detection of chemical cues in murky waters, supported by expanded olfactory regions in braincase reconstructions. The braincase is robust, contributing to balance and equilibrium. Possible salt-excreting glands, inferred from in the antorbital region and prefrontal, would have aided in fully marine lifestyles, analogous to those in other sauropterygians. A prominent pineal , often with a raised rim, may have housed a photosensitive organ for depth gauging or . Variations across subclades highlight adaptive divergence within Pliosauroidea. Rhomaleosaurids possess more slender snouts with relatively larger orbits and moderately sized temporal fenestrae, suited to versatile piscivory and smaller prey in seas, as exemplified by Rhomaleosaurus cramptoni with its spatulate yet elongated premaxillary rostrum. Pliosaurids, by contrast, evolved massive, reinforced crania with expanded temporal regions and vertically oriented quadrates for amplified bite mechanics, evident in the broad, triangular skulls of taxa like Pliosaurus kevani, optimizing for top-predator roles against . These shifts reflect increasing specialization toward and durophagy in later oceans.

Postcranial Skeleton

The postcranial skeleton of pliosauroids is characterized by a robust vertebral column adapted for powerful aquatic propulsion and stability. The neck is notably short, comprising 13 to 30 , which are compact and sub-platycoelous with short centra relative to their height and width, often featuring paired rib facets on the anterior cervicals. This contrasts with the longer necks of plesiosauroids and supports a optimized for rapid maneuvering rather than extended reach. The thoracic region includes approximately 20-25 dorsal vertebrae with robust, mediolaterally compressed that expand the , aiding in regulation through lung volume adjustments. Caudal vertebrae form a sturdy , typically numbering 20-30, with chevron facets for ventral support. , often double-forked and sigmoidal in shape, provide additional ventral reinforcement along the abdominal region. Overall, presacral vertebrae total approximately 35-55, contributing to a compact that enhances hydrodynamic efficiency. The limb girdles and appendages of pliosauroids exhibit advanced modifications for underwater locomotion, with four large, paddle-like flippers serving as primary propulsors. Hyperphalangy is evident, with extra phalanges increasing flipper flexibility and surface area beyond the primitive pentadactyl condition, allowing for undulating motions akin to dynamic flight in modern cetaceans. The and are robust, with expanded proximal and distal ends forming broad articular surfaces; for instance, in , the measures up to 480 mm in length with asymmetrical margins and distinct radial and ulnar facets, while the is slightly longer at around 470-500 mm, featuring a fan-shaped distal expansion for enhanced lift generation during strokes. Fore- and hindflippers are subequal in size, though hindlimbs are often marginally larger, with interlocking phalanges and robust epipodials ( and ) that maintain paddle integrity under thrust. These features indicate a style involving alternating or synchronized beats of all four limbs, generating thrust through hydrodynamic lift rather than drag-based paddling. Body proportions in pliosauroids emphasize a streamlined , with a broad, trunk supported by elongated dorsal ribs that curve posteriorly, minimizing drag while accommodating large internal organs. The overall silhouette is compact, with the presacral region occupying a significant portion of total length (up to 320 cm in some specimens like ), and a short tail providing minor steering assistance. expansion, facilitated by flexible costal elements, likely enabled active control of via inflation, essential for ambushing predators in varied depths. Some taxa show elongated neural spines in the dorsal region, potentially supporting soft-tissue structures resembling low dorsal fins for stability, though direct evidence is limited to inferred musculature attachments. Ontogenetic changes in the pliosauroid skeleton reveal shifts in proportions that refine aquatic adaptations. Juveniles exhibit proportionally longer necks relative to body size, with cervical centra approximately 20% shorter and neural arches unfused, indicating ongoing growth and greater flexibility. As individuals mature, the neck shortens proportionally, centra elongate, and sutures fuse (e.g., neurocentral closure by sub-adult stage), resulting in a more rigid, compact axial skeleton suited to larger body masses and powerful bursts of speed. These transformations, observed in plesiosauroids like Cryptoclidus with parallels in pliosauroids, underscore paedomorphic retention of open sutures in some adults, potentially preserving flexibility.

Classification and Phylogeny

Taxonomy

Pliosauroidea is an extinct clade within the order Plesiosauria, encompassing short-necked marine reptiles that ranged from the Early Jurassic to the Early Late Cretaceous. The superfamily is defined as all plesiosaurians more closely related to Pliosaurus than to Plesiosaurus, forming a monophyletic group supported by synapomorphies such as a short neck with fewer than 20 cervical vertebrae and enlarged temporal fenestrae. Basal forms like those in Neusticosauridae, known from the Late Triassic, are sometimes positioned as outgroups to Pliosauroidea in broader sauropterygian phylogenies, though their exact affinity remains debated. The primary families within Pliosauroidea are Rhomaleosauridae and , with Rhomaleosauridae restricted to the (Hettangian–Toarcian) and characterized by robust skulls, elongated snouts, and four-flipper locomotion adapted for ambush predation. Key genera in Rhomaleosauridae include (type species R. cramptoni, holotype NHMUK PV R.8230, a partial skeleton from the Lower Lias of , ) and Macroplata (type species M. tenuiceps, holotype NHMUK PV R.5830, a skull and vertebrae from the Lower Lias of , ). , the dominant family from the (Hettangian–Tithonian) through the Early Late Cretaceous (Aptian–Turonian), features massive skulls with conical teeth and reduced neck vertebrae, representing macropredatory forms up to 12 meters in length. Within Pliosauridae, notable genera from the Middle to Late Jurassic include Liopleurodon (type species L. ferox, holotype NHMUK PV M.1675, a mandible from the Oxford Clay of Peterborough, England) and Pliosaurus (type species P. brachyspondylus, neotype CAMSM J.35991, a partial skeleton from the Kimmeridge Clay of Norfolk, England). Attenborosaurus (type species A. conybeari, holotype NHMUK PV R.2850, an articulated skeleton from the Lower Lias of Dorset, England) represents an Early Jurassic member of the family. Cretaceous representatives comprise Brachauchenius (type species B. lucasi, holotype KUVP 485, a partial skeleton from the Carlile Shale of Kansas, USA) and Sachicasaurus (type species S. vitae, holotype UPN-08-PV-001, a partial skeleton from the Paja Formation of Colombia). Recent discoveries include new postcranial material of Liopleurodon ferox from the Callovian of central France (specimen ComCom Thouarsais_Geol.0121, ~3.2 m long axial skeleton), confirming its validity and extending its range southward, and additional large pliosaurid remains from the late Valanginian of Colombia, assigned to Sachicasaurus and highlighting persistent diversity in low-latitude settings. Taxonomic revisions have resolved several junior synonyms, particularly within Pliosaurus, where Liopleurodon macromerus is synonymized with P. macromerus based on shared dental morphology (e.g., 70–80 cranial teeth, 8–9 symphyseal pairs) and retroarticular process shape. Ketchum and Benson (2010) established the modern framework by integrating 66 taxa and 178 characters, validating four Pliosaurus species (P. brachyspondylus, P. macromerus, P. brachydeirus, P. funkei) while invalidating others like P. andrewsi due to non-diagnostic traits. Post-2023 updates, including the French Liopleurodon specimen, reinforce paedomorphic traits (e.g., open neurocentral sutures in adults) without altering familial boundaries.

Evolutionary Relationships

Pliosauroidea is defined as a stem-based clade comprising all plesiosaurians more closely related to Pliosaurus than to Plesiosaurus, rendering it the sister group to Plesiosauroidea within the broader clade Plesiosauria. This positioning places Pliosauroidea within Plesiosauria, part of the larger group Pistosauria encompassing advanced sauropterygians, based on cladistic analyses of morphological characters such as vertebral counts and cranial proportions. Key synapomorphies of Pliosauroidea include a reduced cervical vertebral count resulting in a short and an elevated head-to-body size ratio, which distinguish it from long-necked plesiosauroids and facilitate enhanced predatory capabilities. These features are supported by character matrices in comprehensive phylogenetic datasets, initially outlined in Benson et al. (2012) with 66 taxa and 178 characters, and refined in subsequent updates that incorporate new specimens like the 2024 French remains, which affirm the clade's through shared cranial robusticity and dental morphology. Within Pliosauroidea, Rhomaleosauridae occupies a basal position, characterized by early Jurassic forms with moderately large skulls and transitional postcranial features bridging pistosauroids and derived pliosauroids. Pliosauridae forms a monophyletic subclade, with internal branching that groups advanced taxa such as Pliosaurus and Brachauchenius in a derived lineage exhibiting gigantism and specialized bite mechanics. Evidence for paedomorphosis appears in several lineages, particularly in Liopleurodon and related pliosaurids, where retention of juvenile traits like unfused cervical vertebrae in adults suggests heterochronic evolution influencing body proportions and maturity assessment. Phylogenetic controversies persist regarding the inclusion of Cretaceous short-necked forms, with historically allied to but now frequently excluded and placed within based on analyses emphasizing convergences in cranial elongation rather than shared ancestry. Recent discoveries, such as the 2025 Colombian brachauchenine pliosaurid specimens from the upper , further challenge prior views by enhancing documented pliosauroid diversity and supporting a prolonged of into the early .

Evolutionary History

Origins

The origins of Pliosauroidea trace back to Late Triassic precursors within the broader clade Pistosauria, a group of marine sauropterygians that exhibited transitional features between terrestrial reptiles and fully aquatic forms, such as elongated bodies and paddle-like limbs. Pistosaurus, from the of dated to approximately 240 million years ago (Ma), represents a key pistosaurian with a mix of plesiosaur-like axial elongation and more primitive skeletal traits, setting the stage for plesiosaurian evolution. The first unequivocal records of true pliosauroids appear in the Early Jurassic Hettangian stage (around 201–199 Ma), shortly after the end-Triassic mass extinction. Genera such as Stratesaurus taylori and Avalonnectes arturi, known from deposits in the , exemplify basal pliosauroid morphology from the lowermost of . These forms mark the initial emergence of Pliosauroidea as a distinct lineage within Plesiosauria, diverging from plesiosauroid relatives around 200 Ma based on stratigraphic and phylogenetic evidence. Pliosauroids evolved from long-necked sauropterygian ancestors, with early taxa retaining relatively elongated cervical regions compared to later short-necked giants, facilitating a gradual shift toward more predatory adaptations. This transition occurred amid the initial diversification of Plesiosauria in the epicontinental seas of , particularly the shallow shelf environments of the proto-North Atlantic, where post-extinction ecological niches allowed for rapid colonization. In the environmental context of recovering oceans following the end-Triassic extinction event (approximately 201.3 Ma), which decimated marine faunas including many pre-existing reptiles, pliosauroids adapted to warmer, oxygen-rich shelf seas with increased productivity. Their rise paralleled the diversification of other marine predators, such as neoichthyosaurs, which also filled apex roles in these ecosystems, while early fishes began radiating as potential prey in the same intervals. Basal pliosauroid diversity in the Early Jurassic was characterized by small-bodied forms, typically 3–5 meters in length, contrasting with the larger sizes of later representatives. Macroplata tenuiceps, from the Sinemurian of Somerset, England, exemplifies this with an estimated length of about 3 meters and a slender build suited to piscivory in coastal habitats. This early radiation featured high generic diversity but low morphological disparity, reflecting opportunistic exploitation of vacant niches in the post-extinction marine realm.

Diversification and Decline

Pliosauroids underwent a marked diversification in the Mid-Jurassic, with a notable radiation during the and Oxfordian stages, exemplified by the appearance of giant predators such as in European marine environments. This interval witnessed an explosive increase in morphological disparity, particularly in dental adaptations, as pliosaurids achieved their peak variation in tooth morphology with minimal turnover between taxa. Body sizes escalated to over 10 meters, a trend linked to the abundance of larger prey items like teleosaurids and other marine reptiles, allowing these short-necked plesiosaurs to occupy niches across shallow epicontinental seas. During the , pliosauroids maintained a presence but with diminished overall diversity, extending into the stage primarily through forms like in the of . Evidence from the Southern Hemisphere, including the from Colombia's Paja Formation, underscores their persistence in low-latitude settings well into the , representing one of the most complete large-bodied specimens from the region and indicating adaptive resilience in tropical marine habitats. Pliosauroid dispersal was enabled by connectivity through the Tethys Sea, facilitating faunal exchange from European to Mexican assemblages during the , where diversity reached its zenith in the with an estimated 12–20 valid genera across the clade. This global migration supported their role as dominant macropredators until the mid-Cretaceous, when intensified competition from radiations of mosasaurs and , combined with widespread ocean anoxic events and thermal stress, precipitated a sharp decline. The group ultimately went extinct around 90 million years ago, shortly after the Cenomanian-Turonian boundary.

Paleobiology

Diet and Feeding

Pliosauroids employed a puncture-and-tear feeding , utilizing their robust skulls and serrated, conical teeth to seize and dismember prey through powerful post-symphyseal bites rather than shaking or twisting motions. This method was supported by exceptionally high bite forces, estimated at 9.6 to 48 kN in large forms such as those comparable to kevani, enabling them to crush bone and puncture tough hides. Jaw anatomy, with a short, strong rostrum and large adductor muscles, optimized these mechanics for rapid, forceful closure on struggling prey. The prey spectrum of pliosauroids varied with body size, reflecting their role as versatile macropredators in marine ecosystems. Smaller pliosauroids primarily consumed and cephalopods, while gigantic forms targeted larger vertebrates including ichthyosaurs, , and substantial , with evidence also suggesting opportunistic scavenging of carcasses. Prey items could reach up to half the length of the predator, indicating a capacity to tackle sizable targets in open marine environments. Direct evidence of diet derives from preserved stomach contents and associated coprolites, which contain cephalopod hooklets, remains, and bones, confirming a carnivorous regime focused on and vertebrates. Stable isotope analyses of δ¹⁵N in bones further position pliosauroids at high trophic levels (4 or above), consistent with their apex predatory status among marine s. Ontogenetic shifts in diet are inferred from growth-related changes in cranial robusticity and tooth morphology, with juveniles likely piscivorous and focused on softer prey like , transitioning to macropredatory habits targeting larger vertebrates in adulthood. This progression aligns with increasing body size and biomechanical capabilities during maturation.

Locomotion and Sensory Capabilities

Pliosauroids propelled themselves through the water using a four-flipper system that mimicked underwater flight, with the foreflippers providing the primary through powerful, alternating strokes similar to those of modern sea turtles or . The hindflippers contributed secondary and stability, generating up to 60% more and 40% higher when synchronized with the foreflippers in a coordinated, motion. Their short tails likely served mainly as rudders for steering during maneuvers, rather than significant propulsive elements. Flipper articulations in fossils, such as those of , indicate high flexibility at the shoulder and hip joints, enabling rapid adjustments for agile hunting in three dimensions. Estimated swimming speeds for pliosauroids were modest for cruising but capable of bursts for prey pursuit; optimal sustained speeds ranged from 0.4 to 0.5 m/s, with maximum sustained speeds up to 2.5 m/s, based on energetic models and hydrodynamic simulations. These velocities align with comparisons to modern analogs like leatherback turtles, which use similar flipper-driven locomotion for efficient long-distance travel. Trackways attributable to plesiosaurs are exceedingly rare, but biomechanical analyses of flipper morphology support a maneuverable style suited to ambushing prey in open marine environments. Pliosauroids likely achieved through lung inflation and ingestion, allowing them to adjust depth without constant swimming effort; computational models show that with lungs partially inflated, even large individuals could become neutrally buoyant at depths of 10-50 m using stones comprising 1-6% of body mass. Fossil evidence of (decompression syndrome) in bones indicates they routinely dove to depths of 100-200 m or more, comparable to modern deep-diving cetaceans. This buoyancy control facilitated prolonged subsurface hunting, with the compact body form enhancing stability during descent and ascent. The sensory capabilities of pliosauroids were adapted for low-visibility aquatic hunting, featuring large eyes with sclerotic rings that suggest enhanced vision in dim or turbid waters. Orbit sizes in taxa like Pliosaurus indicate forward-directed eyes capable of binocular vision for depth perception during close-range strikes. A complex rostral neurovascular system, innervated by the trigeminal nerve, is preserved in Pliosaurus funkei fossils, implying integumentary sensory organs for detecting water movements or weak electric fields from prey, akin to those in crocodilians. These adaptations collectively supported precise prey localization and ambush tactics in diverse marine settings.

Fossil Record

History of Discovery

The discovery of pliosauroid fossils began in the early along the of , where erosion of the cliffs at revealed remains. In 1841, anatomist formally named the genus Pliosaurus based on fragmentary bones, including vertebrae and limb elements, collected from deposits such as the Formation; he interpreted the animal as an intermediate form between long-necked plesiosaurs and crocodilians due to its robust build and short neck. Owen established species such as P. brachyspondylus and P. brachydeirus, marking the initial recognition of these short-necked predators, though early interpretations suffered from limited material and comparisons to contemporary reptiles. Advancements in the refined pliosauroid classification and expanded the known record. In 1943, paleontologist Samuel P. Welles coined the superfamily name Pliosauroidea in a comprehensive revision, distinguishing short-necked forms with large heads from long-necked plesiosaurs based on skeletal proportions and vertebral counts. Subsequent decades saw incremental finds in and , but significant progress occurred in the and with discoveries beyond traditional sites. A notable example is the "Monster of Aramberri," a partial skeleton unearthed in the late from strata in , ; this specimen, estimated at 10-11 meters in total length, represented the first major pliosauroid from and was formally described in as an indeterminate gigantic pliosaurid. Recent decades have yielded landmark specimens, enhancing understanding of pliosauroid diversity and anatomy. In 2009, a massive approximately 2-meter from the Formation near , , was announced, later designated as the of Pliosaurus kevani in 2013; this find highlighted the group's apex predatory role with its robust jaws and conical teeth. A nearly complete 2-meter discovered in 2023 from , Dorset, by local collector Steve Etches, provided one of the best-preserved pliosauroid crania, revealing details of sensory structures and bite mechanics. In 2024, new partial remains from the of western were attributed to Liopleurodon ferox, expanding its known distribution and confirming paedomorphic traits in this early pliosauroid. Methodological innovations have complemented these field discoveries, particularly through non-invasive imaging and expanded global surveys. Computed tomography (CT) scanning has illuminated internal anatomy, such as the complex neurovascular network in the rostrum of Pliosaurus skulls, traced via bifurcating channels linked to teeth and sensory pits, as demonstrated in analyses from the 2010s onward. International expeditions have increased non-European recoveries, including the brachauchenine Sachicasaurus vitae from the Barremian Paja Formation in Colombia, described in 2019 as a 10-meter predator and the most complete South American pliosauroid to date. These efforts, coupled with reanalyses of museum holdings—like ongoing German studies of Jurassic marine reptiles—continue to uncover pliosauroid evolutionary patterns.

Distribution and Major Localities

Fossil occurrences of pliosauroids are predominantly reported from and marine deposits across Laurasian and later Gondwanan landmasses, reflecting their adaptation to shallow epicontinental seas. In , the primary regions include the , , and , where key formations such as the and in have yielded significant material, including isolated vertebrae, teeth, and partial skulls from the . For instance, the Formation, a bituminous deposited in low-oxygen, lagoonal environments, has produced notable specimens like four large indicating a body length of up to 14 meters. Similarly, the Formation in has provided pliosaurid remains, such as a partial referable to a new , highlighting the formation's role in preserving Middle to marine reptiles. In , the of southwestern regions contains early-diverging pliosauroid elements, though often fragmentary, from lagoonal settings. North American records are centered on the Sundance Formation in , particularly the Upper Redwater Member, which dates to the Oxfordian stage of the and has preserved the largest known pliosaurid from the continent, Megalneusaurus rex, based on limb bones and vertebrae from shallow marine deposits. South American localities, emerging later in the fossil record, include the Paja Formation in Colombia's , where Barremian-aged strata near have yielded the near-complete skeleton of vitae, a brachauchenine pliosaurid exceeding 10 meters in length. A more recent discovery from upper beds in the same region describes two new brachauchenine specimens, expanding the pliosauroid presence in northern and underscoring ongoing exploration in these Gondwanan sites. Stratigraphically, pliosauroid fossils span the , with concentrations in the Callovian-Oxfordian and Kimmeridgian , and extend into the , including the Barremian-Aptian Paja Formation, though records from later formations in remain sparse and unconfirmed for pliosauroids. Taphonomic biases favor preservation in low-energy, anoxic lagoonal and nearshore deposits, such as the organic-rich shales of the , which inhibit scavenging and promote rapid burial. Articulated skeletons are rare due to post-mortem in high-energy marine settings, with most finds consisting of isolated bones, teeth, and partial skulls; however, exceptional preservation has improved with recent recoveries, like the nearly complete 2-meter pliosaur skull from Dorset's in 2023, extracted from cliff exposures. Biogeographically, pliosauroids exhibit early dominance in Laurasian realms during the , with abundant records from European and North American shelves connected by the , transitioning to Gondwanan occurrences in the as widened seaways and facilitated southern dispersal. Examples include the Laurasian forms in the Sundance Formation contrasted with Early Gondwanan taxa like in , illustrating a shift influenced by paleoceanographic barriers post-Pangaea breakup.

Notable Taxa

Early Forms

The basal pliosauroids of the represent the initial radiation of short-necked plesiosaurs, emerging as stem-group members that bridged the gap between Late Triassic pistosaurs and more derived pliosauroids. These early forms, primarily from the to stages (approximately 199–174 million years ago), exhibited transitional traits such as relatively slender builds and necks longer than those of later giant pliosauroids, while retaining robust skulls adapted for predation. Unlike the elongated necks of contemporaneous plesiosauroids, these taxa displayed a of features, including moderately large heads with conical teeth suited for grasping soft-bodied prey, reflecting their role in early marine ecosystems as mid-tier predators. Rhomaleosaurus, a key genus from the of the , exemplifies these early pliosauroids with specimens reaching lengths of about 7 meters. Known from sites like and , it featured a broad skull with curved, pointed teeth indicative of a diet focused on and cephalopods, alongside a relatively long neck comprising approximately 27 —longer proportionally than in later pliosauroids. This genus, often classified within Rhomaleosauridae, dominated European seas as an before the turnover. Smaller-bodied taxa like Hauffiosaurus, from the of and the United Kingdom's coast, highlight the diversity of basal forms with lengths estimated at 3–5 meters. These pliosauroids possessed slender rostra, lateral mandibular troughs, and elongated retroarticular processes, suggesting agile swimming and precise feeding mechanics. Fossils from the often preserve soft tissues, such as skin impressions and gastric contents, providing insights into their piscivorous habits and lightweight construction for burst . A reanalysis of German specimens has confirmed Hauffiosaurus as a distinct basal species, underscoring its transitional position. Macroplata, another Sinemurian representative, further illustrates the slender, piscivorous niche of early pliosauroids, with body lengths up to 4.5 meters and needle-like teeth optimized for capturing fish. Recovered from coastal deposits in , its anatomy includes a wide mandibular symphysis and reduced robusticity compared to later forms, emphasizing speed over power in hunting. These taxa collectively signify the stem-pliosauroid phase, facilitating the evolutionary shift toward the larger, more specialized pliosauroids of the while populating post-Triassic marine recoveries.

Giant Forms

The giant forms of Pliosauroidea, belonging to the clade Thalassophonea, dominated marine ecosystems as macropredatory plesiosaurs, often exceeding 10 meters in total length and featuring disproportionately large skulls relative to body size. These apex predators evolved robust dentitions and powerful jaw mechanics adapted for crushing and tearing large prey, such as other marine reptiles and fish, with their rise marking a shift toward top-tier trophic roles from the onward. Among the earliest giant pliosaurs, from the () Formation of exemplifies this trend, with the largest known skull measuring approximately 1.54 meters in length and body estimates reaching 8–9 meters based on vertebral and limb scaling. Its short neck, broad skull, and conical teeth supported ambush predation in shallow epicontinental seas. Late Jurassic representatives, particularly within Pliosaurus, achieved even greater dimensions. Pliosaurus kevani, described from the Formation (Kimmeridgian, ), possesses a 2-meter-long skull with subtrihedral teeth and an estimated body length of 10–12 meters, inferred from skull-to-body ratios in related pliosaurids. Similarly, Pliosaurus funkei from the of exhibits comparable skull proportions (around 2.2 meters estimated) and body sizes exceeding 12 meters, highlighting a peak in pliosauroid during this interval. An unnamed gigantic specimen from the same locality suggests lengths up to 14.4 meters, based on femur and centrum scaling, potentially representing the largest known pliosauroid. Cretaceous giant forms, though less diverse, include Megacephalosaurus eulerti from the Turonian Carlile Shale of Kansas, USA, with skulls up to 1.75 meters long—75% larger than those of contemporaneous Brachauchenius lucasi (1-meter skulls)—and body lengths estimated at 9 meters via limb bone proportions. These late-surviving pliosaurs retained the short-necked, large-headed bauplan but adapted to more open-ocean environments before the group's extinction in the early Late Cretaceous.

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