extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...

biblical The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of ...

Leviathan Leviathan (; he, לִוְיָתָן, ) is a sea serpent noted in theology and mythology. It is referenced in several books of the Hebrew Bible, including Psalms, the Book of Job, the Book of Isaiah, the Book of Amos, and, according to some ...

Herman Melville Herman Melville ( born Melvill; August 1, 1819 – September 28, 1891) was an American novelist, short story writer, and poet of the American Renaissance period. Among his best-known works are '' Moby-Dick'' (1851); '' Typee'' (1846), a ...

Moby-Dick ''Moby-Dick; or, The Whale'' is an 1851 novel by American writer Herman Melville. The book is the sailor Ishmael's narrative of the obsessive quest of Ahab, captain of the whaling ship ''Pequod'', for revenge against Moby Dick, the giant whi ...

Pisco Formation The Pisco Formation is a geologic formation located in Peru, on the southern coastal desert of Ica and Arequipa. The approximately thick formation was deposited in the Pisco Basin, spanning an age from the Middle Miocene up to the Early Pleisto ...

Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = National seal , national_motto = "Firm and Happy f ...

Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Torto ...

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided ...

year A year or annus is the orbital period of a planetary body, for example, the Earth, moving in its orbit around the Sun. Due to the Earth's axial tilt, the course of a year sees the passing of the seasons, marked by change in weather, the h ...

Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...

South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...

Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58macroraptorial sperm whales (or "raptorial sperm whales") and was probably an

apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...

, preying on whales, seals, and so forth. Characteristically of raptorial sperm whales, ''Livyatan'' had functional, enamel-coated teeth on the upper and lower jaws, as well as several features suitable for hunting large prey. ''Livyatan''s total length has been estimated to be about , similar to that of the modern

sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...

(''Physeter macrocephalus''), making it one of the largest predators known to have existed. The teeth of ''Livyatan'' measured , and are the largest biting teeth of any known animal, excluding tusks. It is distinguished from the other raptorial sperm whales by the basin on the skull, spanning the length of the

snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is ...

. The

spermaceti organ The spermaceti organ is an organ present in the heads of toothed whales of the family Physeteroidea, in particular the sperm whale. This organ contains a waxy liquid called spermaceti and is involved in the generation of sound. Description I ...

, contained in that basin is thought to have been used in echolocation and communication, or for ramming prey and other sperm whales. The whale may have interacted with the large extinct shark

Megalodon Megalodon (''Otodus megalodon''), meaning "big tooth", is an extinct species of mackerel shark that lived approximately 23 to 3.6 million years ago (Mya), from the Early Miocene to the Pliocene epochs. It was formerly thought to be a memb ...

(''Otodus megalodon''),

competing Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...

with it for a similar food source. Its extinction was probably caused by a cooling event at the end of the Miocene period causing a reduction in food populations. The geological formation where the whale has been found has also preserved a large assemblage of marine life, such as sharks and

marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...

s.

Taxonomy

Research history

In November 2008, a partially preserved skull, as well as teeth and the lower jaw, belonging to ''L. melvillei'', the

holotype specimen A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

MUSM 1676, were discovered in the coastal desert of Peru in the sediments of the

Pisco Formation The Pisco Formation is a geologic formation located in Peru, on the southern coastal desert of Ica and Arequipa. The approximately thick formation was deposited in the Pisco Basin, spanning an age from the Middle Miocene up to the Early Pleisto ...

, southwest of the city of Ica. Klaas Post, a researcher for the Natural History Museum Rotterdam in the Netherlands, stumbled across them on the final day of a field trip. The fossils were prepared in

Lima Lima ( ; ), originally founded as Ciudad de Los Reyes (City of The Kings) is the capital and the largest city of Peru. It is located in the valleys of the Chillón, Rímac and Lurín Rivers, in the desert zone of the central coastal part of ...

, and are now part of the collection of the

Museum of Natural History, Lima The Natural History Museum (Museo de Historia Natural) in Lima, is Peru's most important museum of natural history. It was established in 1918 and belongs to the National University of San Marcos. History The museum was founded in 1918 by the Fa ...

of

National University of San Marcos The National University of San Marcos ( es, Universidad Nacional Mayor de San Marcos, link=no, UNMSM) is a public research university located in Lima, the capital of Peru. It is considered the most important, recognized and representative educ ...

. The discoverers originally assigned—in July 2010—the English name of the biblical monster, Leviathan, to the whale as ''Leviathan melvillei''. However, the scientific name ''Leviathan'' was also the

junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...

for the

mastodon A mastodon ( 'breast' + 'tooth') is any proboscidean belonging to the extinct genus ''Mammut'' (family Mammutidae). Mastodons inhabited North and Central America during the late Miocene or late Pliocene up to their extinction at the end of th ...

(''Mammut''), so, in August 2010, the authors rectified this situation by coining a new genus name for the whale, ''Livyatan'', from the original

Hebrew Hebrew (; ; ) is a Northwest Semitic language of the Afroasiatic language family. Historically, it is one of the spoken languages of the Israelites and their longest-surviving descendants, the Jews and Samaritans. It was largely preserved ...

name of the monster. The

species name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

''melvillei'' is a reference to

Herman Melville Herman Melville ( born Melvill; August 1, 1819 – September 28, 1891) was an American novelist, short story writer, and poet of the American Renaissance period. Among his best-known works are '' Moby-Dick'' (1851); '' Typee'' (1846), a ...

, author of the book ''

Moby-Dick ''Moby-Dick; or, The Whale'' is an 1851 novel by American writer Herman Melville. The book is the sailor Ishmael's narrative of the obsessive quest of Ahab, captain of the whaling ship ''Pequod'', for revenge against Moby Dick, the giant whi ...

'', which features a gigantic sperm whale as the main

antagonist An antagonist is a character in a story who is presented as the chief foe of the protagonist. Etymology The English word antagonist comes from the Greek ἀνταγωνιστής – ''antagonistēs'', "opponent, competitor, villain, enemy, riv ...

. The first ''Livyatan'' fossils from Peru were initially dated to around 13–12 million years ago (mya) in the

Serravallian The Serravallian is, in the geologic timescale, an List of time periods, age or a stage (stratigraphy), stage in the middle Miocene epoch (geology), Epoch/series (stratigraphy), Series, which spans the time between 13.82 annum, Ma and 11.63 Ma (m ...

Age of the Miocene, but this was revised to 9.9–8.9 mya in the

Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Torto ...

Age of the Miocene. During the late 2010s, fossils of large isolated sperm whale teeth were reported from various Miocene and Pliocene localities along the Southern Hemisphere. These teeth have been identified to be of similar size and shape with that of the ''L. melvillei'' holotype and may be species of ''Livyatan''. However, it is commonplace that authors do not identify such teeth as a conclusive species of ''Livyatan'', instead opting to assign an

open nomenclature Open nomenclature is a vocabulary of partly informal terms and signs in which a taxonomist may express remarks about their own material. This is in contrast to synonymy lists, in which a taxonomist may express remarks on the work of others. Common ...

in which the biological classifications of the specimens are restricted to comparisons or affinities with ''Livyatan''. This is mostly because isolated teeth tend to not be informative enough to be identified at the species level, meaning that there is some undeterminable possibility that they belong to an undescribed close relative of ''Livyatan'' rather than ''Livyatan'' itself. In 2016 in

Beaumaris Bay Beaumaris Bay ( ) is a bay in the east of Port Phillip, Victoria, Australia. Beaumaris Bay commences at the cliffs at Rickett's Point in the south of the suburb of Beaumaris Beaumaris ( ; cy, Biwmares ) is a town and community on the Isl ...

, Australia, a large sperm whale tooth measuring , specimen NMV P16205, was discovered in Pliocene

strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as ei ...

by a local named Murray Orr, and was nicknamed the "Beaumaris sperm whale" or the "giant sperm whale". The tooth was donated to

Museums Victoria Museums Victoria is an organisation which operates three major state-owned museums in Melbourne, Victoria: the Melbourne Museum, the Immigration Museum and Scienceworks Museum. It also manages the Royal Exhibition Building and a storage facil ...

at

Melbourne Melbourne ( ; Boonwurrung/Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a met ...

. Though it has not been given a species designation, the tooth looks similar to those of ''L. melvillei'', indicating it was a close relative. The tooth is dated to around 5mya, and so is younger than the ''L. melvillei'' holotype by around 4or 5million years. In 2018, palaeontologists led by David Sebastian Piazza, while revising the collections of the Bariloche Paleontological Museum and the Municipal Paleontological Museum of Lamarque, uncovered two incomplete sperm whale teeth cataloged as MML 882 and BAR-2601 that were recovered from the Saladar Member of the

Gran Bajo del Gualicho Formation Gran may refer to: People *Grandmother, affectionately known as "gran" *Gran (name) Places * Gran, the historical German name for Esztergom, a city and the primatial metropolitan see of Hungary * Gran, Norway, a municipality in Innlandet count ...

in the Río Negro Province of

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...

, a deposit that dates between around 20-14 mya. The partial teeth measure and in height respectively. Anatomical analyses of the specimens found that much of their characteristics are identical to ''L. melvillei'' except in width, in which the diameter of both teeth are smaller. Because of this, along with only isolated teeth being available, the palaeontologists chose to assign an open nomenclature, identifying both specimens as ''aff. Livyatan sp.'' In 2019, palaeontologist Romala Govender reported the discovery of two large sperm whale teeth from Pliocene deposits near the Hondeklip Bay village of

Namaqualand Namaqualand (khoekhoe: "Nama-kwa" meaning Nama Khoe people's land) is an arid region of Namibia and South Africa, extending along the west coast over and covering a total area of . It is divided by the lower course of the Orange River into ...

in

South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...

. The pair of teeth, which are stored in the

Iziko South African Museum The Iziko South African Museum is a South African national museum located in Cape Town. The museum was founded in 1825, the first in the country. It has been on its present site in the Company's Garden since 1897. The museum houses important A ...

and cataloged as SAM-PQHB-433 and SAM-PQHB-1519, measure and in height respectively, the latter having its crown missing. Both teeth have open pulp cavities, indicating that both whales were young. The teeth are very similar in shape and size to the mandibular teeth of the ''L. melvillei'' holotype, and were identified as ''cf. Livyatan''. Like the Beaumaris specimen, the South African teeth are dated to around 5mya.

Phylogeny

''Livyatan'' was part of a fossil

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

of hyper-predatory

sperm whales The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the Genus (biology), genus ''Physeter'' and one of three extant species in the s ...

commonly known as macroraptorial sperm whales, or raptorial sperm whales, alongside the extinct whales ''

Brygmophyseter ''Brygmophyseter'', known as the biting sperm whale, is an extinct genus of toothed whale in the sperm whale family with one species, ''B. shigensis''. When it was first described in 1994, the species was placed in the genus ''Scaldicetus'' b ...

'', ''

Acrophyseter ''Acrophyseter'' is a genus of extinct sperm whales that lived in the Late Miocene off the coast of Peru comprising two species: ''A. deinodon'' and ''A. robustus''. It is part of a group of macroraptorial sperm whales which all shared several ...

'', and ''

Zygophyseter ''Zygophyseter varolai'' is an extinct sperm whale that lived during the Tortonian age of the Late Miocene 11.2 to 7.6 million years ago. It is known from a single specimen from the Pietra Leccese Formation in Italy. It was a member of a stem ...

''. This group is known for having large, functional teeth in both the upper and lower jaws, which were used in capturing large prey, and had an enamel coating. Conversely, the modern

sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...

(''Physeter macrocephalus'') lacks teeth in the upper jaw, and the ability to use its teeth to catch prey. ''Livyatan'' belongs to a different lineage in respect to the other raptorial sperm whales, and the size increase and the development of the

spermaceti organ The spermaceti organ is an organ present in the heads of toothed whales of the family Physeteroidea, in particular the sperm whale. This organ contains a waxy liquid called spermaceti and is involved in the generation of sound. Description I ...

, an organ that is characteristic of sperm whales, are thought to have evolved independently from other raptorial sperm whales. The large teeth of the raptorial sperm whales either evolved once in the group with a basilosaurid-like common ancestor, or independently in ''Livyatan''. The large

temporal fossa The temporal fossa is a fossa (shallow depression) on the side of the skull bounded by the temporal lines and terminating below the level of the zygomatic arch. Boundaries * Medial: frontal bone, parietal bone, temporal bone, and sphenoid bon ...

in the skull of raptorial sperm whales is thought to a

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

feature, that is, a trait inherited from a common ancestor. Since the teeth of

foetal A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...

modern sperm whales (''Physeter macrocephalus'') have enamel on them before being coated with

cementum Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament.Illustrated Dental Embryology, Histology, a ...

, it is thought that the enamel is also an ancient characteristic ( basal). The appearance of raptorial sperm whales in the fossil record coincides with the diversification of

baleen whale Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...

s in the

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

, implying that they evolved specifically to exploit baleen whales. It has also been suggested that the raptorial sperm whales should be placed into the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

Hoplocetinae, alongside the genera '' Diaphorocetus'', ''

Idiorophus ''Idiorophus'' is a genus of toothed whales in the family Physeteridae. Fossils have been found in the Colhuehuapian Gaiman Formation of Argentina and the Libano Sandstone in Italy. The teeth of ''Idiorophus'' were similar in size to those of ...

'', ''

Scaldicetus ''Scaldicetus'' is an extinct genus of highly predatory macroraptorial sperm whale. Although widely used for a number of extinct physeterids with primitive dental morphology consisting of enameled teeth, ''Scaldicetus'' as generally recognized ...

'', and '' Hoplocetus'', which are known from the Miocene to the lower

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58wastebasket taxa for non-diagnostic material of stem physeteroids. This subfamily is characterized by their robust and enamel-coated teeth. The cladogram below is modified from Lambert et al. (2017) and represents the phylogenetic relationships between ''Livyatan'' and other sperm whales, with genera identified as macroraptorial sperm whales in bold.

Description

The body length of ''Livyatan'' is unknown since only the holotype skull is preserved. Lambert and colleagues estimated the body length of ''Livyatan'' using ''Zygophyseter'' and modern

sperm whales The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the Genus (biology), genus ''Physeter'' and one of three extant species in the s ...

as a guide''.'' The authors opted to use the relationship between the bizygomatic width (distance between the opposite

zygomatic process The zygomatic processes are three processes (protrusions) from other bones of the skull which each articulate with the zygomatic bone. The three processes are: * Zygomatic process of frontal bone from the frontal bone * Zygomatic process of ma ...

es) of the skull and body length because of the variable

rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...

length in modern sperm whales and the rostrum of ''Livyatan'' being proportionally shorter. Doing so produced length estimates of when using the modern sperm whale and when using ''Zygophyseter.'' It has been estimated to weigh based on the length estimate of . By comparison, the modern sperm whale length measures on average for females and for males, with some males reaching up to long. The large size was probably an

anti-predator adaptation Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avo ...

, and allowed it to feed on larger prey. ''Livyatan'' is the largest fossil sperm whale discovered, and was also one of the biggest-known predators, having the largest bite of any

tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...

.

Skull

The holotype skull of ''Livyatan'' was about long. Like other raptorial sperm whales, ''Livyatan'' had a wide gap in between the

temporal fossa The temporal fossa is a fossa (shallow depression) on the side of the skull bounded by the temporal lines and terminating below the level of the zygomatic arch. Boundaries * Medial: frontal bone, parietal bone, temporal bone, and sphenoid bon ...

e on the sides of the skull and the zygomatic processes on the front of the skull, indicating a large space for holding strong

temporal muscle In anatomy, the temporalis muscle, also known as the temporal muscle, is one of the muscles of mastication (chewing). It is a broad, fan-shaped convergent muscle on each side of the head that fills the temporal fossa, superior to the zygomatic a ...

s, which are the most powerful muscles between the skull and the jaw. The

snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is ...

was robust, thick, and relatively short, which allowed it to clamp down harder and better handle struggling prey. The left and right

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...

e on the snout probably did not intersect at the tip of the snout, though the premaxillae took up most of the front end of the snout. Unlike in the modern sperm whale, the premaxillae reached the sides of the snout. The upper jaw was thick, especially midway through the snout. The snout was asymmetrical, with the right

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...

in the upper jaw becoming slightly convex towards the back of the snout, and the left maxilla becoming slightly concave towards the back of the snout. The

vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...

reached the tip of the snout, and was slightly concave, decreasing in thickness from the back to the front. A sudden thickening in the middle-left side of the vomer may indicate the location of the nose plug muscles. Each

mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...

in the lower jaw was higher than it was wide, with a larger gap in between the two than in the modern sperm whale. The

mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral halves ...

which connects the two halves of the mandibles in the middle of the lower jaw was unfused. The

condyloid process The condyloid process or condylar process is the process on the human and other mammalian species' mandibles that ends in a condyle, the mandibular condyle. It is thicker than the coronoid process of the mandible and consists of two portions: the c ...

, which connects the lower jaw to the skull, was located near the bottom of the mandible like other sperm whales.

Teeth

Unlike the modern sperm whale, ''Livyatan'' had functional teeth in both jaws. The wearing on the teeth indicates that the teeth sheared past each other while biting down, meaning it could bite off large portions of flesh from its prey. Also, the teeth were deeply embedded into the gums and could interlock, which were adaptations to holding struggling prey. None of the teeth of the holotype were complete, and none of the back teeth were well-preserved. The lower jaw contained 22 teeth, and the upper jaw contained 18 teeth. Unlike other sperm whales with functional teeth in the upper jaw, none of the tooth roots were entirely present in the

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...

portion of the snout, being at least partially in the maxilla. Consequently, its tooth count was lower than those sperm whales, and, aside from the modern

dwarf Dwarf or dwarves may refer to: Common uses *Dwarf (folklore), a being from Germanic mythology and folklore * Dwarf, a person or animal with dwarfism Arts, entertainment, and media Fictional entities * Dwarf (''Dungeons & Dragons''), a humanoid ...

(''Kogia sima'') and

pygmy In anthropology, pygmy peoples are ethnic groups whose average height is unusually short. The term pygmyism is used to describe the phenotype of endemic short stature (as opposed to disproportionate dwarfism occurring in isolated cases in a pop ...

(''K. breviceps'') sperm whales, it had the lowest tooth count in the lower jaw of any sperm whale. The most robust teeth in ''Livyatan'' were the fourth, fifth, and sixth teeth in each side of the jaw. The well-preserved teeth all had a height greater than , and the largest teeth of the holotype were the second and third on the left lower jaw, which were calculated to be around high. The first right tooth was the smallest at around . The Beaumaris sperm whale tooth measured around in length, and is the largest fossil tooth discovered in Australia. These teeth are thought to be among the largest of any known animal, excluding

tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...

s. Some of the lower teeth have been shown to contain a facet for when the jaws close, which may have been used to properly fit the largest teeth inside the jaw. In the front teeth, the tooth diameter decreased towards the base. This was the opposite for the back teeth, and the biggest diameters for these teeth were around in the lower jaw. All teeth featured a rapid shortening of the diameter towards the tip of the tooth, which were probably in part due to wearing throughout their lifetimes. The curvature of the teeth decreased from front to back, and the lower teeth were more curved at the tips than the upper teeth. The front teeth projected forward at a 45° angle, and, as in other sperm whales, cementum was probably added onto the teeth throughout the animal's lifetime. Livyatan skull

All tooth sockets were cylindrical and single-rooted. The tooth sockets increased in size from the first to the fourth and then decreased, the fourth being the largest at around in diameter in the upper jaws, which is the largest of any known whale species. The tooth sockets were smaller in the lower jaw than they were in the upper jaw, and they were circular in shape, except for the front sockets which were more ovular.

Basin

The fossil skull of ''Livyatan'' had a curved basin, known as the supracranial basin, which was deep and wide. Unlike other raptorial sperm whales, but much like in the modern sperm whale, the basin spanned the entire length of the snout, causing the entire skull to be concave on the top rather than creating a snout as seen in ''

Zygophyseter ''Zygophyseter varolai'' is an extinct sperm whale that lived during the Tortonian age of the Late Miocene 11.2 to 7.6 million years ago. It is known from a single specimen from the Pietra Leccese Formation in Italy. It was a member of a stem ...

'' and ''

Acrophyseter ''Acrophyseter'' is a genus of extinct sperm whales that lived in the Late Miocene off the coast of Peru comprising two species: ''A. deinodon'' and ''A. robustus''. It is part of a group of macroraptorial sperm whales which all shared several ...

''. The supracranial basin was the deepest and widest over the

braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...

, and, unlike other raptorial sperm whales, it did not overhang the

eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...

. It was defined by high walls on the sides. The antorbital notches, which are usually slit-like notches on the sides of the skull right before the snout, were inside the basin. A slanting crest on the temporal fossa directed towards the back of the skull separated the snout from the rest of the skull, and was defined by a groove starting at the antorbital processes on the cheekbones. The basin had two

foramina In anatomy and osteology, a foramen (;Entry "foramen"
in

in the front, as opposed to the modern sperm whale which has one foramen on the maxilla, and to the modern dwarf and pygmy sperm whales which have several in the basin. The suture in the basin between the maxilla and the

forehead In human anatomy, the forehead is an area of the head bounded by three features, two of the skull and one of the scalp. The top of the forehead is marked by the hairline, the edge of the area where hair on the scalp grows. The bottom of the fore ...

had an interlocking pattern.

Palaeobiology

Hunting

''Livyatan'' was an

apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...

, and probably had a profound impact on the structuring of Miocene marine communities. Using its large and deeply rooted teeth, it is likely to have hunted large prey near the surface, its diet probably consisting mainly of medium-sized baleen whales ranging from in length. It probably also preyed upon sharks, seals, dolphins, and other large marine vertebrates, occupying a

niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...

similar to the modern

killer whale The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only extant species in the genus ''Orcinus'' and is recognizable by its black-and-white pa ...

(''Orcinus orca''). It was contemporaneous with and occupied the same region as the otodontid shark '' O. megalodon'', which was likely also an apex predator, implying

competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...

over their similar food sources. It is assumed that the hunting tactics of ''Livyatan'' for hunting whales were similar to that of the modern killer whale, pursuing prey to wear it out, and then drowning it. Modern killer whales work in groups to isolate and kill whales, but, given its size, ''Livyatan'' may have been able to hunt alone. Isotopic analysis of enamel from a tooth from Chile enamel revealed that this individual likely operated at latitudes south of 40°S. Isotopic analyses of contemporary baleen whales in the same formation show that this ''Livyatan'' was not commonly feeding on them, indicating it probably did not exclusively eat large prey, though it may have targeted baleen whales from higher latitudes.

Spermaceti organ

Sperm whale head anatomy (transverse + sagittal)

The supracranial basin in its head suggests that ''Livyatan'' had a large

spermaceti organ The spermaceti organ is an organ present in the heads of toothed whales of the family Physeteroidea, in particular the sperm whale. This organ contains a waxy liquid called spermaceti and is involved in the generation of sound. Description I ...

, a series of

oil An oil is any nonpolar chemical substance that is composed primarily of hydrocarbons and is hydrophobic (does not mix with water) & lipophilic (mixes with other oils). Oils are usually flammable and surface active. Most oils are unsaturated ...

and

wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to giv ...

reservoirs separated by

connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...

. The uses for the spermaceti organ in ''Livyatan'' are unknown. Much like in the modern sperm whale, it could have been used in the process of

biosonar Echolocation, also called bio sonar, is a biological sonar used by several animal species. Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these ...

to generate sound for locating prey. It is possible that it was also used as a means of acoustic displays, such as for communication purposes between individuals. It may have been used for acoustic stunning, which would have caused the bodily functions of a target animal to shut down from exposure to the intense sounds. Another theory says that the enlarged forehead caused by the presence of the spermaceti organ is used in all sperm whales between males fighting for females during mating season by head-butting each other, including ''Livyatan'' and the modern sperm whale. It may have also been used to ram into prey; if this is the case, in support of this, there have been two reports of modern sperm whales attacking whaling vessels by ramming into them, and the organ is disproportionally larger in male modern sperm whales. An alternate theory is that sperm whales, including ''Livyatan'', can alter the temperature of the wax in the organ to aid in buoyancy. Lowering the temperature increases the density to have it act as a weight for deep-sea diving, and raising the temperature decreases the density to have it pull the whale to the surface.

Palaeoecology

Fossils conclusively identified as ''L. melvillei'' have been found in Peru and Chile. However, additional isolated large sperm whale teeth from other locations including Australia, Argentina, and South Africa have been identified as a species or possible close relative of ''Livyatan''. On the basis of these fossils, it was likely that the distribution of ''Livyatan'' was widespread but restricted to the Southern Hemisphere. The warmer waters around the equator have been known to be a climatic barrier for numerous cetaceans since Neogene times, and a possible hypothesis is that ''Livyatan'' may have been among the cetaceans unable to cross the equatorial barrier. However, due to a poor fossil record of ''Livyatan'', collecting bias is another possible explanation. The holotype of ''L. melvillei'' is from the

Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Torto ...

stage of the Upper

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

9.9–8.9 mya in the Pisco Formation of Peru, which is known for its well-preserved assemblage of marine vertebrates. Among the baleen whales found, the most common was an undescribed species of

cetotheriid Cetotheriidae is a family of baleen whales (parvorder Mysticeti). The family is known to have existed from the Late Oligocene to the Early Pleistocene before going extinct. Although some phylogenetic studies conducted by recovered the living ...

whale measuring around , and most of the other baleen whales found were roughly the same size. Toothed whale remains found consist of beaked whales (such as ''

Messapicetus ''Messapicetus'' is an extinct genus of beaked whale from the Late Miocene. It currently holds two species, ''M. longirostris'' from the Tortonian of Italy and ''M. gregarius'' from the Pisco Formation of Peru. However, a third unnamed species is ...

gregarius''), ancient pontoporiids (such as '' Brachydelphis mazeasi''),

oceanic dolphin Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as the ...

s, and the raptorial sperm whale ''Acrophyseter''. All seal remains found represent the earless seals. Also found were large sea turtles such as ''

Pacifichelys ''Pacifichelys'' is an extinct genus of sea turtle from the Middle Miocene of Peru (Pisco Formation) and California ( Temblor Formation).seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the orde ...

es in this area. Partial bones of crocodiles were discovered. Of the seabirds, fragmentary bones of

cormorant Phalacrocoracidae is a family of approximately 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the IOC adopted a consensus taxonomy of seven ge ...

s and

petrels Petrels are tube-nosed seabirds in the bird order Procellariiformes. Description The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group (all except the albatross f ...

were discovered, as well as two species of

boobies A booby is a seabird in the genus ''Sula'', part of the family Sulidae. Boobies are closely related to the gannets (''Morus''), which were formerly included in ''Sula''. Systematics and evolution The genus ''Sula'' was introduced by the Fren ...

. The remains of many

cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...

were discovered in this formation, including more than 3,500 shark teeth, which mainly belonged to the ground sharks, such as

requiem shark Requiem sharks are sharks of the family Carcharhinidae in the order Carcharhiniformes. They are migratory, live-bearing sharks of warm seas (sometimes of brackish or fresh water) and include such species as the tiger shark, bull shark, le ...

s and

hammerhead shark The hammerhead sharks are a group of sharks that form the family Sphyrnidae, so named for the unusual and distinctive structure of their heads, which are flattened and laterally extended into a "hammer" shape called a cephalofoil. Most hammerhe ...

s. To a lesser extent,

mackerel sharks The Lamniformes (, from Greek ''lamna'' "fish of prey") are an order of sharks commonly known as mackerel sharks (which may also refer specifically to the family Lamnidae). It includes some of the most familiar species of sharks, such as the gr ...

were also found, such as

white sharks The great white shark (''Carcharodon carcharias''), also known as the white shark, white pointer, or simply great white, is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is nota ...

,

sand shark Sand sharks, also known as sand tiger sharks, gray nurse sharks or ragged tooth sharks, are Lamniformes, mackerel sharks of the family (biology), family Odontaspididae. They are found worldwide in temperate and tropical waters. The three specie ...

s, and

Otodontidae Otodontidae is an extinct family of sharks belonging to the order Lamniformes. Its members have been described as megatoothed sharks. They lived from the Early Cretaceous to the Pliocene, and included genera such as ''Carcharocles'' and ''Otod ...

. Many shark teeth were associated with the extinct broad-tooth mako (''Cosmopolitodus/Carcharodon hastalis'') and megalodon, and the teeth of these two sharks were found near whale and seal remains.

Eagle ray The eagle rays are a group of cartilaginous fishes in the family Myliobatidae, consisting mostly of large species living in the open ocean rather than on the sea bottom. Eagle rays feed on mollusks and crustaceans, crushing their shells with the ...

s,

sawfish Sawfish, also known as carpenter sharks, are a family of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined with sharp transverse teeth, arranged in a way that resembles a saw. They are among the largest fish wi ...

, and

angelshark The angelsharks are a group of sharks in the genus ''Squatina'' of the family Squatinidae. They commonly inhabit sandy seabeds close to in depth. Many species are now classified as critically endangered by the International Union for Conservat ...

s were other cartilaginous fish found. Most of the

bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...

findings belonged to

tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...

s and

croakers Sciaenidae are a family of fish in the order Acanthuriformes. They are commonly called drums or croakers in reference to the repetitive throbbing or drumming sounds they make. The family consists of about 286 to 298 species in about 66 to 70 gene ...

. ''Livyatan'' and megalodon were likely the apex predators of this area during this time. ''L. melvillei'' is also known from the

Bahía Inglesa Formation Caldera Basin ( es, Cuenca de Caldera) is a sedimentary basin located in the coast of northern Chile west of Copiapó. The basin has a fill of marine sediments of Late Cenozoic age. With a north-south extension of and an east-west width of the ...

of Chile, whose fossiliferous beds are dated between the Tortonian and

Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the first ...

9.03–6.45 mya. Like the Pisco Formation, the Bahía Inglesa Formation famously holds one of the richest marine vertebrate assemblages. Baleen whale remains include ancient

minke whale The minke whale (), or lesser rorqual, is a species complex of baleen whale. The two species of minke whale are the common (or northern) minke whale and the Antarctic (or southern) minke whale. The minke whale was first described by the Danish n ...

s,

grey whale The gray whale (''Eschrichtius robustus''), also known as the grey whale,Britannica Micro.: v. IV, p. 693. gray back whale, Pacific gray whale, Korean gray whale, or California gray whale, is a baleen whale that migrates between feeding and bree ...

s,

bowhead whale The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus ''Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters, ...

s, and cetotheriids. Of the toothed whales, five species of pontoporiids as well as beaked whales,

porpoise Porpoises are a group of fully aquatic marine mammals, all of which are classified under the family Phocoenidae, parvorder Odontoceti (toothed whales). Although similar in appearance to dolphins, they are more closely related to narwhals an ...

s, three other species of sperm whales such as ''cf. Scaldicetus'', and the ''

Odobenocetops ''Odobenocetops'' () is an extinct genus of small toothed whale known from Peru and Chile. Its fossils are found in Neogene-aged marine strata dating from the Tortonian to the Zanclean. It had two tusks, and, in some fossils, one tusk was longer t ...

'' have been yielded. Other marine mammals include the marine sloth ''

Thalassocnus ''Thalassocnus'' is an extinct genus of semiaquatic ground sloths from the Miocene and Pliocene of the Pacific South American coast. It is monotypic within the subfamily Thalassocninae. The five species—''T. antiquus'', ''T. natans'', ''T. lit ...

'' and pinnipeds like '' Acrophoca''. At least 28 different species of sharks have been described, including many extant ground sharks and white sharks as well as extinct species such as the false mako (''Parotodus sp.''), broad-toothed mako, megalodon, and the transitional great white ''

Carcharodon hubbelli ''Carcharodon hubbelli'', also known as Hubbell's white shark, is an extinct species of white shark that evolved between 8 and 5 million years ago during the Late Miocene to Early Pliocene epochs. The shark is a transitional species, showing inter ...

''. Other marine vertebrates include

penguin Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...

s and other seabirds, and species of crocodiles and

ghavial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...

s. The Beaumaris sperm whale was found in the

Beaumaris Bay Beaumaris Bay ( ) is a bay in the east of Port Phillip, Victoria, Australia. Beaumaris Bay commences at the cliffs at Rickett's Point in the south of the suburb of Beaumaris Beaumaris ( ; cy, Biwmares ) is a town and community on the Isl ...

Black Rock Sandstone Formation in Australia near the city of

Melbourne Melbourne ( ; Boonwurrung/Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a met ...

, dating to 5mya in the Pliocene. Beaumaris Bay is one of the most productive marine fossil sites in Australia for marine megafauna. Shark teeth belonging to twenty different species have been discovered there, such as from the

whale shark The whale shark (''Rhincodon typus'') is a slow-moving, filter-feeding carpet shark and the largest known extant fish species. The largest confirmed individual had a length of .McClain CR, Balk MA, Benfield MC, Branch TA, Chen C, Cosgrove J, D ...

(''Rhincodon typus''), the

Port Jackson shark The Port Jackson shark (''Heterodontus portusjacksoni'') is a nocturnal, oviparous (egg laying) type of bullhead shark of the family Heterodontidae, found in the coastal region of southern Australia, including the waters off Port Jackson. It has ...

(''Heterodontus portusjacksoni''), the broad-toothed mako, and megalodon. Some examples of whales found include the ancient humpback whale '' Megaptera miocaena'', the

dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the ...

'' Steno cudmorei'', and the sperm whale '' Physetodon baileyi''. Other large marine animals found include ancient

elephant seal Elephant seals are very large, oceangoing earless seals in the genus ''Mirounga''. Both species, the northern elephant seal (''M. angustirostris'') and the southern elephant seal (''M. leonina''), were hunted to the brink of extinction for oil ...

s,

dugong The dugong (; ''Dugong dugon'') is a marine mammal. It is one of four living species of the order Sirenia, which also includes three species of manatees. It is the only living representative of the once-diverse family Dugongidae; its closest m ...

s, sea turtles, ancient penguins such as ''

Pseudaptenodytes The extinct penguin genus ''Pseudaptenodytes'' contains the type species ''P. macraei''; smaller bones have been assigned to ''P. minor'', although it is not certain whether they are really from a different species or simply of younger individual ...

'', the extinct albatross '' Diomedea thyridata'', and the extinct toothed seabirds of the genus ''

Pelagornis ''Pelagornis'' is a widespread genus of prehistoric pseudotooth birds. These were probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to account for this uncertain ...

''. The South African teeth attributed as ''cf. Livyatan'' are from the Avontuur Member of the

Alexander Bay Formation Alexander is a male given name. The most prominent bearer of the name is Alexander the Great, the king of the Ancient Greek kingdom of Macedonia (ancient kingdom), Macedonia who created one of the largest empires in ancient history. Variants li ...

near the village of Hondeklip Bay, Namaqualand, which is also dated to around 5mya in the Pliocene. The Hondeklip Bay locality enjoys a rich heritage of marine fossils, whose diversity may have been thanks to the initiation of the Benguela Upwelling during the late Miocene, which likely provided large populations of phytoplankton traveling the cold nutrient-rich waters. Cetaceans are the most abundant fauna in the bay, although remains tend to be difficult to conclusively identify. Included are three species of balaenopterids including two undetermined species and one identified as ''cf. Plesiobalaenoptera'', an ancient grey whale (''cf. Eschrichtius sp.''), an undetermined

balaenid Balaenidae () is a Family (biology), family of whales of the parvorder Mysticeti that contains two living genera: the right whales (genus ''Eubalaena''), and in a separate genus, the closely related bowhead whale (genus ''Balaena''). Evolutionar ...

, an unidentified dolphin, and another undetermined species of macroraptorial sperm whale. Other localities of similar age on the South African west coast have also yielded many additional species of balaenopterids and sperm whales as well as ten species of beaked whales. Large sperm whale teeth of up to around ~ in length are common in Hondeklip Bay, indicating a high presence of large sperm whales like ''Livyatan'' in the area. The locality has also a high presence of sharks indicated by a large abundance of shark teeth; however, most of these teeth have not been identified. Megalodon teeth have been found in the bay, and evidence from bite marks in whale bones indicate the additional presence of the

great white shark The great white shark (''Carcharodon carcharias''), also known as the white shark, white pointer, or simply great white, is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is nota ...

,

shortfin mako The shortfin mako shark (; ; ''Isurus oxyrinchus''), also known as the blue pointer or bonito shark, is a large Lamniformes, mackerel shark. It is commonly referred to as the mako shark, as is the longfin mako shark (''Isurus paucus''). The shor ...

, and broad-toothed mako. Other marine fauna known in Hondeklip Bay include pinnipeds such as '' Homiphoca capensis'', bony fish, and rays.

Extinction

''Livyatan''-like sperm whales became extinct by the early Pliocene likely due to a cooling trend causing baleen whales to increase in size and decrease in diversity, becoming coextinct with the smaller whales they fed on. Their extinction also coincides with the emergence of the killer whale as well as large predatory globicephaline dolphins, possibly acting as an additional stressor to their already collapsing niche.

References

External links

'Sea monster' fossil found in Peru desert
by

CNN CNN (Cable News Network) is a multinational cable news channel headquartered in Atlanta, Georgia, U.S. Founded in 1980 by American media proprietor Ted Turner and Reese Schonfeld as a 24-hour cable news channel, and presently owned by ...

Ancient sperm whale's giant head uncovered
from Los Angeles Times (The skull is on display in National History Museum in Lima)
A killer sperm whale
with details on sperm whale evolution ;Videos * * * (a collection of images of discovery) {{Taxonbar, from=Q131759 Miocene mammals of Africa Sperm whales Prehistoric toothed whales Prehistoric cetacean genera Miocene cetaceans Miocene mammals of South America Neogene Peru Fossils of Peru Pisco Formation Fossil taxa described in 2010 Herman Melville