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Carcharodon

Carcharodon is a genus of mackerel sharks (order Lamniformes, family Lamnidae) comprising one extant species, the great white shark (C. carcharias), an iconic apex predator renowned for its size, powerful build, and serrated teeth adapted for tearing flesh.^[1]^[2] The genus name derives from the Greek words karcharos (sharp) and odous (tooth), reflecting the distinctive triangular, saw-edged dentition of its members.^[2] Taxonomically, Carcharodon belongs to the family Lamnidae, which includes other large sharks like makos and porbeagles, with the great white as the only living representative of the genus.^[1]^[3] Fossil records indicate the Lamnidae family originated in the Paleocene or early Eocene epoch, while the genus Carcharodon appeared in the late Oligocene or early Miocene, with several extinct species such as C. hastalis and C. hubbelli documented from Miocene to Pliocene deposits, though their precise phylogenetic placement remains debated among paleontologists, with some proposing separation into genera like Cosmopolitodus.^[2]^[4] The great white shark exhibits a streamlined, spindle-shaped body reaching up to 6 meters in length and 2,000–3,000 kg in weight, with females typically larger than males; its dorsal surface is grayish, contrasting with a white ventral side for countershading camouflage.^[3]^[2] It possesses regional endothermy, maintaining elevated body temperatures via a rete mirabile vascular system, which enhances muscle performance in cooler waters unlike most ectothermic sharks.^[2] The snout is conical and blunt, the dorsal fin triangular, and the caudal fin lunate, all contributing to its hydrodynamic efficiency for bursts of speed up to 56 km/h.^[3] Distributed cosmopolitically in temperate and subtropical seas from 60°N to 60°S, C. carcharias inhabits coastal waters, continental shelves, and offshore areas up to 1,200 meters deep, with key populations off California, South Africa, Australia, and the Mediterranean.^[3]^[1] As a solitary, migratory predator, it preys primarily on marine mammals like seals and sea lions, as well as fish, cetaceans, and carrion, using ambush tactics and powerful jaw bites with estimates up to 18,000 Newtons of force.^[3]^[5] Reproduction is ovoviviparous, with gestation lasting approximately 12-18 months and litters of 2–14 pups born live after intra-uterine cannibalism among siblings.^[3] Evolutionarily, Carcharodon likely descended from broad-toothed mako ancestors (Isurus hastalis) in the Miocene, with transitional fossils like C. hubbelli bridging to modern forms, though links to the extinct giant shark Carcharocles megalodon are contested and generally rejected in favor of separate lineages.^[2]^[4] Conservationally, the species is classified as Vulnerable by the IUCN due to threats from bycatch, habitat degradation, and historical overfishing, with protections under CITES Appendix II and regional bans on targeted fishing.^[2] Despite its fearsome reputation, human attacks are rare, with great white sharks involved in approximately 15-25 unprovoked incidents annually worldwide as of recent data.^[3]^[6]

Taxonomy and Classification

Etymology and History

The genus name Carcharodon derives from the Ancient Greek words karcharos (κάρχαρος), meaning "sharp" or "pointed," and odous (ὀδούς), meaning "tooth," referring to the shark's serrated, triangular teeth.^[2] The name was first proposed by British zoologist Sir Andrew Smith in the work of German anatomists Johannes Peter Müller and Friedrich Gustav Jacob Henle, published in 1839, to classify the great white shark (Carcharodon carcharias).^[7] Prior to this establishment, the great white shark underwent several taxonomic reassignments in the late 18th and early 19th centuries. It was initially described by Carl Linnaeus in 1758 as Squalus carcharias, placing it within the broad genus Squalus for various cartilaginous fishes. Subsequent classifications shifted it to genera such as Carcharias and Isurus, reflecting early uncertainties in distinguishing mackerel shark morphologies based on limited specimens and anatomical studies.^[2] A major historical debate centered on the inclusion of the extinct giant shark Megalodon within Carcharodon, initially named Carcharodon megalodon by Louis Agassiz in his 1843 monograph Recherches sur les Poissons Fossiles, based on similarities in tooth serration and robust form.^[8] This assignment persisted into the mid-20th century, implying a direct lineage with the great white shark, but was challenged by comparative dental analyses revealing distinct evolutionary paths; Megalodon exhibited broader, finer-serrated teeth suited to different prey.^[9] By the 1960s, Edouard Casier proposed reclassifying it to the separate genus Carcharocles to reflect these morphological differences, a view supported by further revisions in the 1990s and 2000s.^[10] More recent phylogenetic studies, including those from 2012 onward, have shifted Megalodon to the genus Otodus within the extinct family Otodontidae, emphasizing its divergence from Carcharodon during the Paleogene.^[9] The genus Carcharodon saw further refinements through the 19th and 20th centuries, with Agassiz's 1843–1844 descriptions contributing key fossil species like Oxyrhina hastalis (later reassigned), which helped delineate Carcharodon's boundaries based on tooth shape and stratigraphy.^[7] Post-1950s synonymies addressed overlaps with extinct forms; for instance, in 1964, L. S. Glikman erected Cosmopolitodus for hastalis (previously under Isurus or Carcharodon), recognizing it as a broad-toothed mako-like ancestor, though later works (e.g., 2012) treated it as a subgenus of Carcharodon due to transitional fossils linking it to C. carcharias. In 2021, Isurus planus was reassigned to Cosmopolitodus planus, making it the second species in the genus, though the taxonomic status remains debated.^[11] These revisions underscore Carcharodon's position within the family Lamnidae, informed by ongoing tooth-based taxonomy.^[7]

Phylogenetic Relationships

The genus Carcharodon is classified within the family Lamnidae (mackerel sharks) and the subfamily Lamninae, a placement supported by shared morphological features such as a fusiform body plan and physiological adaptations including regional endothermy, which enables elevated temperatures in locomotor musculature, cranial regions, and viscera through vascular counter-current heat exchange systems. This endothermic capability distinguishes Lamnidae from most other shark families and is considered a key synapomorphy for the group, facilitating enhanced swimming performance and predatory efficiency in diverse oceanic environments. Cladistic analyses, incorporating both morphological and molecular data, consistently position Carcharodon as the sister genus to Isurus (mako sharks) within Lamnidae, forming a monophyletic clade characterized by similar dentition patterns and body proportions optimized for fast cruising.^[12] Genetic studies utilizing mitochondrial DNA sequences have reinforced this relationship, with bootstrap support values ranging from 85% to 97% for the monophyly of Isurus and Carcharodon in maximum parsimony and likelihood analyses. More recent mitochondrial genome comparisons from the 2020s continue to uphold this sister-group status, highlighting low genetic divergence and shared haplotypes that suggest a relatively recent divergence within the late Paleogene or early Neogene.^[13] Morphological evidence from dentition and skeletal structure further links Carcharodon to ancestral lamniform sharks, such as the Cretaceous genus Cretalamna, through progressive development of serrated tooth margins and a robust, triangular tooth crown morphology adapted for cutting flesh. Cretalamna teeth lack the fine serrations seen in modern Carcharodon but share a similar overall form and jaw architecture, indicating an evolutionary continuum within Lamniformes from Mesozoic stem-group taxa.^[14] Debates persist regarding the monophyly of Carcharodon, particularly concerning the inclusion of fossil species like C. hastalis, with 2010s proposals advocating its separation into the distinct genus Cosmopolitodus based on vertebral column morphology and the absence of serrations in anterior teeth, which align more closely with broad-toothed mako-like ancestors.^[7] These taxonomic revisions, supported by exceptional fossil skeletons from Miocene deposits, suggest C. hastalis represents a transitional form between Isurus-like makos and the serrated-toothed Carcharodon carcharias, challenging the traditional lumping of these taxa and emphasizing the need for integrated morphometric and phylogenetic analyses.^[15]

Physical Characteristics

Anatomy and Morphology

Sharks of the genus Carcharodon possess a cartilaginous endoskeleton typical of elasmobranchs, composed primarily of flexible hyaline cartilage reinforced in the vertebral column by calcified prismatic blocks that form the centra, providing structural support while maintaining lightweight buoyancy.^[16] This calcified cartilage in the vertebrae enhances rigidity without the density of bone, allowing for efficient swimming in the genus's predatory lifestyle.^[17] Sensory adaptations include the ampullae of Lorenzini, clusters of gel-filled electroreceptive pores concentrated on the ventral snout, which detect weak bioelectric fields emitted by prey, enabling precise localization even in turbid or low-visibility conditions.^[18] For eye protection during feeding, Carcharodon species feature a nictitating membrane, a translucent third eyelid that slides horizontally across the cornea to shield it from damage without impairing vision.^[2] As members of the Lamniformes order, Carcharodon sharks exhibit a robust, spindle-shaped body with a conical snout that optimizes hydrodynamics for burst swimming, five gill slits for efficient oxygen extraction, two dorsal fins for stability, an anal fin for maneuverability, and a lunate caudal fin with a prominent lower lobe that generates powerful propulsion via thunniform locomotion. These traits support the genus's role as apex predators in pelagic environments. Dentition consists of large, triangular teeth with serrated edges, adapted for slashing and gripping flesh; in C. carcharias, upper jaw teeth can reach up to 5 cm in height, while the lower jaw features slightly narrower forms for securing prey.^[19] Teeth are continually replaced in a conveyor-belt fashion, with a replacement rate of approximately 1-2 weeks per row, ensuring functional cutting edges throughout the shark's life.^[20] Carcharodon species demonstrate regional endothermy through a rete mirabile, a countercurrent heat exchange network of arterial and venous capillaries in the musculature, viscera, and cranium, which conserves metabolic heat to elevate core temperatures by 10-15°C above ambient water, facilitating sustained activity in cooler oceanic regions.^[21] This physiological innovation, shared among lamnids, enhances metabolic efficiency and sensory performance in the genus.^[22]

Size Variations Across Species

The great white shark, Carcharodon carcharias, exhibits significant size variation, with adults typically averaging 4 to 5 meters in total length (TL), though verified maximum lengths reach up to approximately 6.0 meters based on specimens measured in the 1980s.^[23] Weight estimates for C. carcharias range from 680 to 2,000 kilograms, reflecting pronounced sexual dimorphism where females attain larger sizes than males, often exceeding 1,500 kilograms compared to males around 1,000 kilograms.^[23]^[3] Among extinct species, Carcharodon hubbelli from the early Pliocene reached comparable dimensions to modern great whites, with body length estimates of approximately 4.9 to 5.1 meters derived from partial skeletons including articulated dentitions and vertebrae.^[24] Weight for C. hubbelli is estimated around 1,500 kilograms based on these skeletal remains, suggesting a robust build similar to C. carcharias despite slower growth trajectories.^[25] In contrast, the late Miocene to Pliocene Cosmopolitodus hastalis (formerly classified as Carcharodon hastalis) is estimated at approximately 5 meters TL on average as of 2025, with reliable ranges of 2.7 to 8.2 meters and maximums exceeding 8 meters inferred from tooth scaling in associated dentitions.^[24]^[26] Growth rates in C. carcharias are slow, with maturity reached at 3.5 to 4 meters TL; males typically mature at around 3.6 meters and 26 years of age, while females do so at 4.2 meters and 33 years, determined through von Bertalanffy growth models applied to annual band counts in vertebrae.^[27]^[28] These models, which fit parameters like asymptotic length (L_∞) at 5.4 to 6.1 meters and growth coefficient (k) of 0.06 to 0.20 per year, validate age estimates up to 73 years via bomb radiocarbon assays on vertebral cores.^[29]^[30] For extinct forms like C. hastalis and C. hubbelli, allometric scaling relates tooth crown height (CH) to total body length via regression equations such as TL ≈ 10.2 × CH (with CH in decimeters and TL in meters), allowing size reconstruction from isolated teeth by correlating crown height to jaw width and overall proportions.^[31] This approach highlights that C. hastalis teeth, averaging 2.5 to 7.5 centimeters in height, yield body estimates comparable to or slightly smaller than those of C. carcharias in some cases.^[4]

Habitat and Distribution

Geographic Range

The genus Carcharodon exhibits a broad global distribution, with its sole extant species, C. carcharias, displaying a cosmopolitan pattern across temperate and subtropical marine waters worldwide, spanning approximately 60°N to 60°S latitude.^[3] This species is commonly encountered in coastal regions such as those off California in the northeastern Pacific, the waters surrounding South Africa in the southeastern Atlantic and southwestern Indian Oceans, and the Mediterranean Sea in the northeastern Atlantic.^[32]^[2]^[33] Fossil evidence reveals that extinct species within the genus occupied extensive spatial ranges during the Neogene. C. hastalis, known from the Miocene, has been documented through teeth in marine deposits across both the Atlantic and Pacific basins, including sites in Florida, California, and Chile.^[4]^[34] In contrast, C. hubbelli appears more geographically restricted, with fossils primarily recovered from Pliocene formations in Peru and Chile along the southeastern Pacific coast.^[7]^[35] Modern C. carcharias individuals demonstrate remarkable migratory capabilities, with satellite tagging efforts—initiated in the 1990s—revealing annual travels of up to 20,000 km.^[36]^[37] For instance, sharks tagged off the coast of Mexico have been tracked migrating to Hawaiian waters in the central Pacific.^[38]^[39] Paleontological records indicate post-Pleistocene range contractions for the genus, likely influenced by climatic shifts at the end of the last glacial period, as evidenced by fossil occurrences in regions that have since become predominantly tropical or subtropical.^[40]^[41]

Environmental Preferences

Carcharodon carcharias, the great white shark, primarily inhabits coastal and continental shelf waters at depths of 0 to 200 meters, though individuals have been recorded tolerating depths up to 1,200 meters.^[19]^[42] These sharks exhibit a preference for shallow continental shelf areas, often remaining within the top 50 meters where they spend the majority of their time.^[43] Salinity levels in their preferred habitats typically range from 30 to 35 parts per thousand (ppt), aligning with standard marine conditions, with tagged juveniles showing selection for surface salinities between 31 and 32 ppt.^[44]^[45] Optimal water temperatures for C. carcharias fall between 12 and 24°C, enabling effective foraging and metabolic function within temperate regions.^[19]^[3] This species employs a regional endothermy system via countercurrent heat exchange mechanisms in vascular retia mirabilia, which conserve metabolic heat generated by red swimming muscles to maintain elevated temperatures in the brain, eyes, stomach, and locomotor muscles during dives into colder water layers below the preferred range.^[46]^[47] In terms of substrate associations, great white sharks favor structured environments such as rocky reefs and kelp forests, which provide cover and ambush opportunities in coastal zones.^[48]^[49] They generally avoid hypersaline waters exceeding typical oceanic levels or freshwater intrusions, reflecting their limited tolerance for significant salinity fluctuations beyond 30-35 ppt.^[50] Additionally, C. carcharias shows adaptations to upwelling zones, such as those off the California coast, where nutrient-rich cold waters enhance prey availability and support higher productivity.^[51] These sharks undertake diel vertical migrations, often following thermoclines to access prey layers while optimizing energy use, with deeper excursions during daylight hours into cooler strata below the surface mixed layer.^[43]^[52] Such movements allow them to exploit vertical gradients in temperature and oxygen within their shelf habitats.^[53]

Behavior and Ecology

Feeding and Predatory Strategies

The genus Carcharodon comprises apex predators with specialized carnivorous diets centered on high-energy marine vertebrates. For the extant species C. carcharias, the great white shark, primary prey consists of marine mammals such as pinnipeds (seals and sea lions) and small cetaceans, which provide nutrient-dense blubber accounting for a substantial portion of caloric intake due to its high lipid content.^[54] Secondary prey items include teleost fishes, elasmobranchs, and seabirds, reflecting opportunistic foraging in coastal and offshore environments.^[55] Hunting strategies in C. carcharias emphasize ambush predation, with sharks often approaching prey from below in a stealthy patrol within receptive fields near pinniped colonies, maintaining low-energy swimming speeds of around 1.3 m/s before explosive breaches.^[56] These breaches can propel the shark upward at high velocities, estimated up to 40 km/h in observational studies, enabling surprise attacks that disorient or injure targets before a follow-up consumption phase. A common tactic involves initial "test bites" to evaluate prey quality and energy yield, often followed by a "hit-and-run" approach in the majority of encounters, where the shark retreats to avoid counterattacks or assess viability, with success rates around 50% in early morning hunts near seal haul-outs.^[56] The predatory apparatus of Carcharodon species features a protrusible upper jaw that extends forward during strikes, combined with serrated triangular teeth designed for slashing and gripping. Computer modeling estimates the bite force of a large C. carcharias at approximately 18,000 N, facilitating deep tissue penetration and efficient prey dismemberment. Tooth rake marks preserved on fossil marine mammal bones from Miocene to Pliocene deposits suggest that extinct congeners, such as C. hubbelli, employed analogous slashing and puncturing techniques against large-bodied prey.^[57] As an apex predator, C. carcharias occupies a trophic level of approximately 4.5, exerting top-down control on marine food webs through selective predation on high-trophic-level species.^[58] Juveniles exhibit a dietary shift toward lower-trophic elasmobranchs and teleosts, as revealed by stomach content analyses from South African specimens in the 1970s and 1980s, which documented over 60% fish remains in sharks under 3 m in length compared to predominantly mammalian diets in adults.^[59] White sharks (Carcharodon carcharias) exhibit ovoviviparous reproduction, characterized by internal fertilization and the development of embryos within the mother without a placental connection. Gestation periods last 12-18 months, after which females give birth to litters of 2-14 pups, with litter size varying based on maternal body size.^[32]^[60] During intrauterine development, embryos engage in oophagy, consuming unfertilized eggs produced by the mother and occasionally siblings to sustain growth, which limits litter sizes but results in larger, better-nourished offspring at birth.^[32]^[61] Females follow a biennial reproductive cycle, breeding approximately every two years, while males mature earlier, reaching sexual maturity at smaller sizes (around 3.5-4.0 m total length) compared to females (4.5-5.0 m). Courtship rituals, including parallel swimming alongside potential mates and subtle bites to the flanks or fins, have been documented in mixed-sex aggregations off Guadalupe Island, Mexico, where adults converge seasonally.^[32]^[62]^[38] Socially, white sharks are predominantly solitary predators, but they form transient, loose aggregations of up to 20 individuals at pinniped colonies, such as those off California, primarily to exploit concentrated prey resources. Acoustic tagging studies from the 2010s, involving over 100 individuals, reveal these gatherings lack stable hierarchies or preferential associations, with sharks exhibiting independent movement patterns despite spatial overlap.^[63]^[64]^[65] Females can live up to 70 years, as estimated through radiocarbon dating of ocular structures like eye lenses, which record long-term growth layers; this extended lifespan, combined with low reproductive output, heightens their susceptibility to population declines from human impacts.^[66]^[67]

Species

Extant Species

The genus Carcharodon is represented by a single extant species, Carcharodon carcharias, commonly known as the great white shark, the type species of the genus.^[68] First described by Carl Linnaeus in 1758 in the 10th edition of Systema Naturae, it is known for its large size and distinctive dentition.^[68]^[69] Population estimates for C. carcharias in the northeastern Pacific vary, with a 2013 National Marine Fisheries Service (NMFS) status review suggesting approximately 3,000 individuals, though recent studies (as of 2023) estimate around 300 in central California aggregations and indicate an overall increasing trend.^[32] Globally, the number of mature individuals is thought to be under 3,500, reflecting limited survey coverage. The species has been assessed as Vulnerable by the International Union for Conservation of Nature (IUCN) since 1996, primarily due to ongoing mortality from bycatch in commercial fisheries targeting tuna and billfish. Genomic analyses in the 2020s have identified distinct genetic clusters among C. carcharias subpopulations, separating those in the Indo-Pacific (including around Australia, South Africa, and New Zealand) from those in the North Atlantic, with a third lineage in the North Pacific. These clusters exhibit low gene flow, evidenced by significant divergence in mitochondrial DNA despite relative uniformity in nuclear genomes, suggesting long-term isolation dating back approximately 100,000–200,000 years. Recent acoustic monitoring efforts have confirmed nursery areas for C. carcharias off New Zealand, particularly in Tauranga Harbour, where tagged juveniles and young-of-the-year sharks have been detected in shallow coastal waters since 2023, highlighting active pupping sites essential for recruitment.^[70]

Extinct Species

The taxonomic placement of these extinct species within Carcharodon is debated, with some paleontologists assigning C. hastalis to the genus Cosmopolitodus. Cosmopolitodus hastalis, often referred to as the broad-tooth mako, is an extinct species known from the Miocene to Pliocene epochs, spanning approximately 15 to 3.6 million years ago (Ma).^[4]^[7]^[71] Its teeth feature broad, triangular crowns that are labiolingually flattened and typically unserrated or with very fine serrations, distinguishing them from the coarser serrations seen in the modern great white shark (C. carcharias). These dental characteristics suggest a predatory adaptation for slicing through softer prey, such as fish and smaller marine vertebrates. Fossil evidence indicates that C. hastalis attained an estimated body length of around 4 meters, based on bite mark analyses on cetacean bones. The species was widespread, with remains documented across various marine deposits, including those in the ancient Paratethys Sea, as well as in the Pacific Basin regions like Peru, Chile, California, and Japan. Another extinct species, Carcharodon hubbelli, represents a transitional form and was described from the Upper Neogene Pisco Formation in Peru. Dated to the late Miocene around 6 to 8 Ma (with some records extending into the early Pliocene at approximately 5 to 3.5 Ma), this species exhibits intermediate tooth morphology, including weakly serrated triangular crowns that bridge features of C. hastalis and C. carcharias. The holotype, consisting of a complete set of jaws with 222 teeth and 45 vertebrae, supports an estimated total length of 4.9 to 5.1 meters. Named in 2012 after researcher Gordon Hubbell, C. hubbelli provides key evidence for the evolutionary progression within the genus, highlighting gradual changes in dentition suited for increasingly robust prey. Fossils are primarily known from the Sud Sacaco locality in the Pisco Formation, underscoring its Pacific coastal distribution.^[7]^[72] The validity of Carcharodon caifassii remains questioned, with limited fossil material attributed to this late Miocene species (approximately 11 to 5 Ma). Originally described from European deposits, it has been reassessed in recent studies as a junior synonym of C. carcharias. No distinct morphological or temporal separation has been conclusively established from the modern species, leading to its de-emphasis in modern classifications.^[73] The extinction of C. hastalis occurred around 3.6 Ma during the early Pliocene, coinciding with a decline in available prey resources, including marine mammals, amid broader ecological shifts such as cooling oceans and increased competition from emerging serrated-toothed lamnids. Fossil records show no temporal overlap between C. hastalis and the modern C. carcharias until the late Pliocene, approximately 3 Ma, marking a clear faunal turnover within the genus.^[7]^[74]

Evolutionary History

Fossil Record Overview

The fossil record of the genus Carcharodon is predominantly composed of isolated teeth, as the cartilaginous skeletons of chondrichthyans rarely preserve due to rapid decay in marine environments.^[75] This reliance on dental remains has yielded abundant Carcharodon teeth from Miocene sites worldwide, providing key insights into the genus's paleoecology and distribution.^[76] The direct stratigraphic range of Carcharodon fossils spans from the Late Oligocene to Early Miocene (approximately 28–16 Ma, with some debated records in the Burdigalian stage) to the present, with peak diversity occurring during the Tortonian stage (approximately 11–7 Ma) in the Late Miocene.^[76] ^[77] Notable global localities include the Sharktooth Hill Bonebed in California, which preserves abundant Carcharodon teeth within Middle Miocene marine sediments, and the Yorktown Formation in Virginia, yielding specimens from Upper Miocene to Lower Pliocene deposits.^[78] Preservation of Carcharodon teeth faces challenges such as bioerosion by microbial and boring organisms, which can degrade surface structures, and recrystallization of the enameloid layer, altering its original mineral composition over time.^[79] Since the 2010s, advancements like micro-computed tomography (micro-CT) scanning have enabled non-destructive 3D modeling of tooth morphology, revealing internal details obscured by taphonomic alterations.^[80] Fossils of Carcharodon often co-occur with remains of baleen whales and sirenians in Miocene marine deposits, suggesting the genus inhabited ecosystems rich in marine megafauna that served as potential prey or associates.^[81]^[82]

Transitional Forms and Lineages

The evolutionary lineage of the genus Carcharodon originates from the Paleocene shark Isurolamna inflata, a piscivorous species with smooth, narrow teeth that represents an early ancestor in the lamniform sharks, though the direct fossil record of Carcharodon sensu stricto begins in the Miocene. The inclusion of species like C. hastalis in the genus Carcharodon is debated, with some researchers assigning it to the separate genus Cosmopolitodus.^[83] Transitional forms between Isurolamna and the Miocene Carcharodon hastalis exhibit progressive increases in tooth robusticity, particularly around the Eocene-Oligocene boundary, where crowns became broader and roots more calcified, adapting to harder prey items.^[84] In the Miocene, the genus underwent a significant radiation, with C. hastalis emerging as the basal species, characterized by triangular teeth initially lacking serrations but capable of inflicting damage on marine mammals, as demonstrated by fossil teeth embedded in cetacean vertebrae from ~14 million years ago.^[84] Serrations subsequently evolved in later C. hastalis variants to enhance cutting efficiency against mammalian prey, marking a shift toward specialized macropredation. Carcharodon hubbelli, dated to approximately 6–8 million years ago in the late Miocene Pisco Formation of Peru, serves as a critical transitional taxon, featuring intermediate tooth morphologies with developing serrations and robusticity between C. hastalis and the modern C. carcharias.^[7] Key adaptive shifts in the Carcharodon lineage transitioned from piscivory in Paleogene ancestors like Isurolamna to macropredation on marine mammals by the Miocene, evidenced by changes in tooth crown shape and implied jaw mechanics for higher bite forces on tougher tissues.^[84] The Messinian salinity crisis (~5.96–5.33 million years ago), followed by the catastrophic Zanclean reflooding, drove Mediterranean speciation by enabling Indo-Pacific colonists to enter via the Atlantic, establishing a distinct C. carcharias population in the Pliocene.^[85] The modern C. carcharias lineage diverged around 4–5 million years ago, coinciding with the appearance of fully serrated teeth in the fossil record. Genetic analyses, including proxies from historical and subfossil samples, reveal population bottlenecks during Pleistocene glaciations, contributing to reduced genetic diversity in contemporary populations.^[86]

Conservation and Human Interactions

Current Threats

The great white shark (Carcharodon carcharias) faces significant anthropogenic threats that contribute to population declines across its range. Bycatch in gillnets and longlines remains a primary concern, particularly in Mediterranean and Australian fisheries where driftnet and longline operations inadvertently capture large predators.^[87] These fisheries target species like tuna and swordfish, but great whites suffer high post-release mortality due to their slow recovery from hooking or entanglement.^[32] Finning for the global shark fin trade and trophy hunting for jaws and teeth exacerbate direct mortality, driven by demand in markets for traditional medicine and collectibles. Although international trade in great white parts is now regulated, illegal poaching persists in regions with weak enforcement.^[88] Habitat degradation further compounds vulnerabilities, as overfishing of key prey like seals and sea lions reduces food availability and forces sharks into riskier foraging behaviors near human activities.^[89] Coastal pollution, including heavy metals and organochlorines, accumulates in nearshore nurseries where juveniles reside, with contaminants transferred from mothers to pups via yolk sacs, impairing early development and survival.^[90] Climate change amplifies these pressures through ocean warming, prompting poleward range shifts in great white distributions as thermal tolerances are exceeded in equatorial and temperate zones.^[91]

Protection Measures

The great white shark (Carcharodon carcharias), the sole extant species in the genus Carcharodon, benefits from several international conservation measures aimed at regulating trade and protecting migratory populations. It has been listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) since 2002, which requires permits for international trade to ensure it does not threaten the species' survival.^[92] Additionally, since 2014, it has been included in Appendix I and II of the Convention on the Conservation of Migratory Species of Wild Animals (CMS), obligating signatory states to conserve migratory populations and prohibit capture except under exceptional circumstances. These listings facilitate coordinated global efforts to monitor and restrict exploitation, particularly given the shark's wide-ranging migrations across ocean basins. In November 2025, Mexico implemented a ban on the capture and retention of several threatened Atlantic sharks, including great whites, to address bycatch and direct fishing pressures.^[93] At the national level, targeted bans have provided full protection in key habitats. South Africa pioneered such measures in 1991 by prohibiting the intentional killing, landing, or sale of great white sharks, setting a precedent for domestic safeguards.^[94] Australia followed in 1999, declaring the species protected under its Environment Protection and Biodiversity Conservation Act and listing it as vulnerable, which bans targeted fishing and trade.^[95] In California, protections took effect in 1994 through state regulations that prohibit the take, possession, or sale of white sharks in state waters, significantly reducing directed fisheries.^[96] More recently, the European Union implemented a ban on the retention, transshipment, landing, or sale of great white sharks in 2018 via General Fisheries Commission for the Mediterranean (GFCM) Recommendation GFCM/41/2018/5, applicable to Mediterranean waters under EU jurisdiction.^[97] Monitoring and management programs further support these protections by addressing incidental interactions. The Food and Agriculture Organization (FAO) of the United Nations promotes shark finning prohibitions through its 1999 International Plan of Action for the Conservation and Management of Sharks (IPOA-Sharks), which over 50 countries have adopted as national plans; these require full utilization of shark carcasses and apply to species like the great white shark to curb fin trade. In the Atlantic, the International Commission for the Conservation of Atlantic Tunas (ICCAT) introduced measures in 2023 under Recommendation 23-12 to limit incidental catch, mandating the live release of great white sharks encountered in tuna and swordfish fisheries with minimal harm.^[98] Evidence of recovery underscores the effectiveness of these interventions. In the Northeast Pacific, photo-identification surveys conducted at aggregation sites off California and Baja California have documented a population rebound of approximately 300% since the 1990s, with adult estimates rising from fewer than 100 individuals to 300-500 by the 2020s, attributed to reduced fishing pressure post-protection.^[62] To mitigate potential impacts from human activities, ecotourism regulations—such as those in South Africa requiring licensed operators, limited vessel numbers, and no-chumming protocols during cage-diving—help minimize behavioral disturbances to sharks while generating conservation funding. The species is currently assessed as Vulnerable globally by the International Union for Conservation of Nature (IUCN), though regional populations such as those in the Mediterranean are classified as Critically Endangered.^[99]

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Megalodon Shark Evolution - Research and Science - by Lutz Andres
Hence, the Megatooth Shark would have to be renamed Otodus megalodon and the Great White Shark Isurus carcharias. At present, however, Carcharocles megalodon is ...<|separator|>
[9]
[PDF] ORIGIN OF THE WHITE SHARK CARCHARODON (LAMNIFORMES
As such, the genus Carcharodon (Smith in Müller and Henle, 1838) is the senior synonym of Cosmopolitodus (Glikman, 1964) the proper genus name for C. hastalis.Missing: etymology | Show results with:etymology
[10]
Phylogeny of lamniform sharks (Chondrichthyes - J-Stage
The exact relationships among Carcharodon and the two Isurus species are uncertain, but they constitute a monophyletic group. The group is sister to the clade.
[11]
Complete Mitochondrial DNA Genome of Nine Species of Sharks ...
This study sequenced the mitochondrial DNA of nine sharks and rays, confirming the monophyly of sharks and rays and their phylogenetic placement.
[12]
Carcharodon - an overview | ScienceDirect Topics
Carcharodon is defined as a monotypic genus within the family Carcharodontidae, originating in the Paleocene and evolving primarily from fish feeders to marine ...Missing: credible | Show results with:credible
[13]
(PDF) A well preserved skeleton of the fossil shark Cosmopolitodus ...
Jan 18, 2017 · Here we report on a partial skeleton of a lamniform shark, including portions of the visceral arches and the anterior segment of the vertebral column.
[14]
[PDF] Shark centra microanatomy and mineral density variation
Elasmobranch (sharks, rays, skates) skeletons consist of cartilage, and the cartilage of shark spine, where the mechanical demands are extreme, is reinforced by ...
[15]
Incremental growth and diagenesis of skeletal parts of the lamnoid ...
Shark skeletons are composed of hyaline cartilage, a vertebrate tissue not prone to fossilization. In a notable exception, the vertebral centra of most sharks, ...
[16]
Great White Shark - Smithsonian Ocean
A series of pores on the shark's snout are filled with cells called the Ampullae of Lorenzini that can feel the power and direction of electrical currents.
[17]
A shark's infinite regeneration of teeth
Feb 17, 2020 · Sharks do not rely on two sets of teeth – they have an endless supply of teeth, with a dentition that regenerates constantly throughout life.
[18]
Carcharodon carcharias, Great white shark - FishBase
Elasmobranchii (sharks and rays) > Lamniformes (Mackerel sharks) > Lamnidae (Mackerel sharks or white shark) Etymology: Carcharodon: carchar-, presumably ...<|control11|><|separator|>
[19]
Ontogenetic Influences on Endothermy in the Great White Shark ...
May 12, 2016 · The great white shark is endothermic. Ecophysiology in relation to endothermy is largely unknown yet may disclose life history ecology.
[20]
Endothermic Sharks - Sharks4kids
Feb 1, 2019 · By using the Rete mirabile, the family Lamnidae can achieve actions their ectothermic cousins can't. By raising their body temperature, they are ...
[21]
Great White Shark Facts and Information | United Parks & Resorts
Lamniformes; Family: Lamnidae; Genus Species: Carcharodon carcharias. Fast Facts. Description: Large-sized, predatory shark with rows of serrated, triangular ...Missing: list authoritative source - -
[22]
Estimating lamniform body size - Palaeontologia Electronica
Mar 1, 2021 · These reconstructions all included a medially-oriented upper intermediate tooth, based on an assumed evolutionary relationship with Carcharodon ...
[23]
Exceptional Preservation of the White Shark Carcharodon ...
Aug 10, 2025 · In these deposits, Cosmopolitodus hastalis is represented by isolated teeth that are commonly found in the early Miocene beds of the Chilcatay ...
[24]
Age and growth of the white shark, Carcharodon carcharias, in the ...
Jan 6, 2015 · Based on the current study and these minimum size estimates, age at maturity is 26 and 33 years for male and female white sharks respectively.
[25]
(PDF) Age and growth determination of the white shark ...
The smallest mature male had 8 GRs (293 cm PCL); there were no mature females. Von Bertalanffy parameters for the combined sexes were L(∞)=544 cm PCL, k=0.065/ ...
[26]
[PDF] Age and growth of the white shark, Carcharodon carcharias, in the ...
Jan 6, 2015 · These age estimates, validated up to 44 years, were used to develop a growth curve for the species, which was best described using the Schnute ...
[27]
Age, growth and genetic status of the white shark (Carcharodon ...
The von Bertalanffy growth parameters were estimated at L∞ = 455 cm TL, k = 0.196 year–1 and t0 = –1.92 years for males and L∞ = 607 cm TL, k = 0.159 year–1 ...
[28]
The relationship between the tooth size and total body length in the ...
To estimate the TL of each Otodus megalodon individual, they used Shimada's (2003) linear regression equations showing the relationships between the tooth crown ...
[29]
White Shark | NOAA Fisheries
Scientific Classification. Kingdom, Animalia. Phylum, Chordata. Class, Chondrichthyes. Order, Lamniformes. Family, Lamnidae. Genus, Carcharodon. Species ...Missing: authoritative | Show results with:authoritative
[30]
The Deep Past of the White Shark, Carcharodon carcharias, in the ...
Oct 20, 2023 · The deep past of C. carcharias in the Mediterranean is witnessed by a rather conspicuous, mostly Pliocene fossil record.
[31]
https://www.researchgate.net/publication/288676146_The_relationship_between_the_tooth_size_and_total_body_length_in_the_white_shark_Carcharodon_carcharias_Lamniformes_Lamnidae
[32]
Carcharodon hastalis - Chile - The Fossil Forum
Oct 7, 2020 · I have here an alleged Carcharodon hastalis from Chile. It is 1.85" long and 1.12" wide. After receiving the tooth, it looked a lot more ...
[33]
The effects of pop-up satellite archival tags (PSATs) on the ...
Pop-up satellite archival tags (PSATs) have been used since the 1990s to document the movements and post-release survival of numerous fish species.
[34]
Great White Shark swims 12,400 miles, shocks scientists - Mongabay
Oct 6, 2005 · Nicole's complete journey of more than more than 12,400 miles (more than 20,000 kilometers) is by far the longest distance traveled by any shark ...
[35]
Two-year migration of adult female white sharks (Carcharodon ...
Apr 4, 2013 · Satellite tagging programs have provided detailed information about the migratory patterns of northeastern Pacific white sharks, ...
[36]
Satellite tagging maps the secret migration of white sharks - Phys.org
Apr 3, 2013 · Female white sharks were found to follow a two-year migration pattern with four distinct phases. Firstly the pregnant females left Guadalupe ...
[37]
[PDF] Carcharodon carcharias) off southeastern
shark that has a cosmopolitan distribution throughout temperate seas and oceans and occa- sionally penetrates tropical zones (Fergusson,. 1996; Compagno ...<|control11|><|separator|>
[38]
Oldest record of the Great White Shark (Lamnidae, Carcharodon
Fossil and archaeological records of Carcharodon carcharias in southern South America. Frenguelli, 1920, Frenguelli, 1922 identified a broken tooth as ...
[39]
Species: Carcharodon carcharias, Great white Shark, White Shark
Habitat: coastal, inshore to oceanic. Depth: 0 to 1280 m. Circumglobal, temperate; Alaska to the Gulf of California; unconfirmed repots in the Galapagos ...
[40]
Movements, behavior and habitat preferences of juvenile white ...
May 24, 2025 · The white shark Carcharodon carcharias is an apex predator with a cosmopolitan distribution in temperate and tropical waters of both ...Missing: zones | Show results with:zones
[41]
(PDF) Carcharodon carcharias-White Shark. The IUCN Red List of ...
Background Understanding white shark (Carcharodon carcharias) habitat use in coastal areas adjacent to large cities, is an important step when formulating ...
[42]
[PDF] Three-Dimensional Movements and Habitat Selection of Young ...
Mar 22, 2021 · Overall, selection ratio results illustrated a preference for sea surface salinities between 31.0 and. 32.0 ppt, with neutral responses to ...
[43]
Adaptations of the Great White Shark
The Great White has such heat exchangers around its brain, stomach, and swimming muscles, enabling it to function effectively in waters too cold for most other ...
[44]
Why Are Great White Sharks Warm-Blooded When ... - A-Z Animals
Feb 24, 2025 · Great white sharks are partially warm-blooded since the rete mirabile only circulates heat to certain parts of the body and in certain environments.
[45]
[PDF] The influence of kelp density on white shark presence within the ...
demonstrated that highly dense kelp forests (i.e. ≥1 stalk of kelp per ... shark (Carcharodon carcharias) during post hoc video analysis. The selected ...
[46]
[PDF] Shallow Rocky Reefs and Kelp Forests - Tom W Bell
such as the great white shark (Carcharodon carcharias), Califor- nia ... Like terres- trial forests, kelp forests modify the physical environment by.
[47]
Three-Dimensional Movements and Habitat Selection of Young ...
Mar 21, 2021 · Recent research has confirmed a white shark (Carcharodon carcharias) summer nursery off Long Island, New York; however, habitat characterization ...
[48]
[PDF] Status Review of the - the NOAA Institutional Repository
Ecosystem as a biologically rich and dynamic ocean environment with a coastal upwelling ... Carcharodon carcharias. Pages 85-90 in M. L. Domeier, editor. Global ...
[49]
[PDF] Thermocline influence on vertical behaviour of marine predators
We found a strong support that thermocline drives the movement of four species of top predatory fish (sharks and tunas). Results showed that turning points ...
[50]
[PDF] Broad-Scale Vertical and Horizontal Behavior of the White Shark ...
This thesis studies the broad-scale vertical and horizontal behavior of white sharks in the Western North Atlantic, including vertical and regional migrations.
[51]
[PDF] Great White Shark (Carcharodon carcharias) - NOAA
Jun 20, 2012 · WildEarth Guardians also requests that NMFS designate critical habitat for the species in U. S. waters. ... IUCN Red List: Carcharodon carcharias.
[52]
Diet Composition and Nutritional Niche Breadth Variability in ...
Jun 6, 2020 · We have provided the first detailed description of the diet of juvenile white sharks in eastern Australia, revealing the importance of both ...
[53]
(PDF) The hunting strategy of white shark (Carcharodon carcharias ...
Our aim was to describe the behavior and interactions among white sharks hunting seals near a seal colony.
[54]
Tooth marks of the Great White Shark from a Pliocene ... - Paleopolis
Jul 24, 2024 · white shark tooth marks are relatively rare compared to the abundance of cetacean finds from the Pliocene epoch, during which Carcharodon ...Missing: strategies rake
[55]
Standardized diet compositions and trophic levels of sharks
The second highest. TL (4.5) corresponded to the great white shark,. Carcharhodon carcharias (Lamniformes). Table 3. Trophic levels of sharks by order (in bold) ...
[56]
Size-based analysis of diet and trophic position of the white shark ...
Previous dietary-related studies on white sharks have shown the species to be an opportunistic carnivore having a diverse diet consisting of fin fishes, small ...
[57]
[PDF] White shark Species Status Assessment - NY.gov
Jan 12, 2024 · Seasonal distribution and historic trends in abundance of white sharks,. Carcharodon carcharias, in the western North Atlantic Ocean. PloS one, ...
[58]
[PDF] Reproductive Modes in Elasmobranchs
reproductive modes typically only result in a few large offspring per litter. Oophagy is unique to just a few species of sharks, such as white sharks.Missing: ovoviviparous | Show results with:ovoviviparous
[59]
White Shark Information - California Department of Fish and Wildlife
Oct 3, 2025 · White sharks, also called great white sharks (Carcharodon carcharias), are one of the ocean's primary predators, and fascinating creatures to many.
[60]
Social dynamics and individual hunting tactics of white sharks ... - NIH
Mar 23, 2022 · We used biologging (activity, cameras and telemetry receivers) sensors to measure social associations and simultaneous behaviour, in white sharks.
[61]
Marine Ecology Progress Series 644:215
Jun 25, 2020 · Analyzing 412 trips to sea by 8 seals and more than 25 000 acoustic detections from 23 individual white sharks, we observed seasonally ...
[62]
[PDF] Co-occurrence of White Sharks (Carcharodon carcharias) along ...
May 2, 2025 · The network analysis of acoustic telemetry data in this study provides insight into the social dynamics of white sharks off the coast of Cape ...
[63]
https://pmc.ncbi.nlm.nih.gov/articles/PMC8941395/
[64]
Radiocarbon dating suggests white sharks can live 70 years and ...
Jan 8, 2014 · Adult white sharks, also known as great whites, may live far longer than previously thought, according to a new study that used radiocarbon dating.Missing: eye lens
[65]
https://repository.lib.umassd.edu/view/pdfCoverPage?instCode=01MA_DM_INST&filePid=13158727600001301&download=true
[66]
Carcharodon carcharias, Great white shark - FishBase
Short description Identification keys | Morphology | Morphometrics · Dorsal spines (total): 0; Anal spines: 0. A huge, spindle-shaped shark with conspicuous ...
[67]
Spying on our great white sharks | Earth Sciences New Zealand
The sharks were tagged with acoustic and popup tags, and filmed underwater for photo-identification purposes. The tags and photos will allow scientists to ...Missing: nursery pupping<|control11|><|separator|>
[68]
Cosmopolitodus Hastalis | Encyclopedia MDPI
Oct 28, 2022 · Carcharodon carcharias ... "A morphometric approach for addressing toothbased species delimitation in fossil mako sharks, Isurus (Elasmobranchii: ...
[69]
New ancient shark species gives insight into origin of great white
Nov 4, 2012 · hubbelli, also known as Hubbell's white shark, was based on a complete set of jaws with 222 teeth intact and 45 vertebrae. The species was named ...
[70]
Carcharodon hastalis | Shark-References
Carcharodon hastalis. (Agassiz, 1838). Classification: Elasmobranchii Lamniformes Lamnidae. Reference of the original description. Agassiz ...
[71]
The Deep Past of the White Shark, Carcharodon carcharias ... - MDPI
... length of at least 615 cm, that is, larger than the largest verified extant specimen of C. carcharias worldwide. Other teeth in the same size range as the ...
[72]
Decoding the dynamics of dental distributions: insights from shark ...
Jun 29, 2022 · Shark species in the fossil record are largely identified by their tooth morphology [23] due to the poor preservation of cartilaginous skeletons ...
[73]
A Review of the Paleobiology of Some Neogene Sharks and ... - MDPI
It had an Early Miocene East Pacific–Central West Atlantic center of origin [161]. The present-day distributional pattern of C. brachyurus is the product of ...
[74]
Return to Sharktooth Hill, Kern County, California
For over 150 years, amateurs and professionals alike have collected Miocene marine fossils from Sharktooth Hill and associated outcrops throughout Kern County, ...Missing: record | Show results with:record
[75]
Fossil microbial shark tooth decay documents in situ metabolism of ...
Dec 1, 2020 · We document here for the first time in situ bacteria preservation in tooth enameloid, one of the hardest mineralized tissues developed by animals.Missing: challenges | Show results with:challenges
[76]
Great white work: Scientists renew the study of shark teeth
Aug 8, 2012 · The microanatomical study of shark teeth has been dormant for decades, but the scientists are renewing it using the latest imaging technology.Missing: 2010s | Show results with:2010s<|separator|>
[77]
Miocene | BVMNH - Buena Vista Museum of Natural History
On display on BVM's main floor, you will find large specimens from Miocene baleen whales, sperm whales, the large sea lion allodesmus and the largest shark to ...
[78]
Repeated mass strandings of Miocene marine mammals from ...
Apr 22, 2014 · Here, we describe an unusually dense accumulation of fossil marine vertebrates from Cerro Ballena, a Late Miocene locality in Atacama Region of ...
[79]
Fossil History of the White Shark
Therefore, Carcharodon and Isurus both descended from Isurolamna inflata and many smooth-edged fossil teeth originally named Isurus are in fact part of the ...Missing: transitional hubbelli
[80]
[PDF] New light on the trophic ecology of Carcharodon hastalis from teeth ...
Jun 16, 2025 · Recent isotopic analyses of the teeth of the extinct lamnid Carcharodon hastalis showed that it fed at a comparable trophic level as was the ...
[81]
Is There Genetic Evidence of a Recent Population Bottleneck in ...
We show that contemporary northwest Atlantic white sharks are genetically distinct from other populations and comprise a demographically distinct unit (pairwise ...
[82]
Global shark fishing mortality still rising despite ... - Science
Jan 11, 2024 · Here we estimate that total fishing mortality increased from at least 76 to 80 million sharks between 2012 and 2019, ~25 million of which were threatened ...
[83]
Fifty Years After 'Jaws,' We've Learned a Lot About Great Whites
Feb 2, 2024 · In the wake of Jaws, shark trophy hunting increased. Sharks were killed for sport, not for food. In a scene forecasting what would follow up ...
[84]
Shark Finning: Sharks Turned Prey | Smithsonian Ocean
This is the process of slicing off a shark's fin and discarding the rest of the still-living body, often by dumping it back into the ocean.
[85]
Are we starving great whites to death? - Conservation Action Trust
Dec 19, 2020 · “Great whites' numbers likely plummeted because of diminished food resources, paired with decades of poaching, bycatch and KwaZulu shark board ...
[86]
Impacts of Ocean Pollution on Sharks and Rays
Similarly, high organochlorine levels have been found in young White Shark pups, presumed to be transferred from their mother's tissue. ... coastal nursery areas ...
[87]
Study finds climate change shrinks and shifts juvenile white shark ...
Feb 9, 2021 · Researchers conclude the northward range shift demonstrates the young sharks are being subjected to a loss of suitable thermal habitat, meaning ...Missing: great models 2050
[88]
Great white shark - CITES
Great white shark ; Scientific name: Carcharodon carcharias ; Distribution: Worldwide distribution ; CITES listing: Appendix II ; Photo: © Rachel Powell ; < Species ...Missing: 2002 | Show results with:2002
[89]
[PDF] p. 1 AC20 Inf. 23 Convention on International Trade in Endangered ...
The Great White Shark is taken as bycatch in fisheries that use longlines, hook-and-line, fixed bottom gillnets, fish traps, herring weirs, and trammelnets, ...
[90]
Great white shark research - Save Our Sharks
The Great White Shark was declared a protected species in 1996 by the NSW Government. In 1999 their threatened species status was elevated to “Vulnerable” to ...Missing: 1991 California 1994
[91]
[PDF] 4 White Shark, Carcharodon carcharias - CA.gov
It has been speculated that females give birth to live litters of 4 to 14 pups, but this is based on a very limited number of pregnant females that have been ...
[92]
New Evidence for the Contemporary Presence of Juvenile White ...
The retention, transshipment, landing, sale, or display of white sharks is strictly prohibited (GFCM 2018, 2021). ... Occurrence of great white shark ...
[93]
https://www.forbes.com/sites/melissacristinamarquez/2025/11/15/mexico-finally-bans-fishing-of-threatened-atlantic-sharks/