Acrophyseter Deinodon
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''Acrophyseter'' is a
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 ...
of extinct
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 '' Physeter'' and one of three extant species in the sperm whale fam ...
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 whale Macroraptorial sperm whales were highly predatory whales of the sperm whale superfamily (Physeteroidea) of the Miocene epoch that hunted large marine mammals, including other whales, using their large teeth. They consist of five genera: ''Acrophy ...
s which all shared several features for the purpose of hunting large prey, such as deeply-rooted and thick teeth. ''Acrophyseter'' measured , making it the smallest raptorial sperm whale. Because of its short pointed snout, and its strong curved front teeth, it probably fed on the large marine vertebrates of its time, such as seals and other whales.


Discovery

The
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 ...
''Acrophyseter'' houses two species. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
, ''A. deinodon'', was discovered in the Sud-Sacaco
locality Locality may refer to: * Locality (association), an association of community regeneration organizations in England * Locality (linguistics) * Locality (settlement) * Suburbs and localities (Australia), in which a locality is a geographic subdivis ...
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 ...
in Peru, dating back to 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 ...
Messinian stages of the
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
around 8.5–6.7
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago) ...
(mya); 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 several ...
, MNHN SAS 1626, represents a mature individual and consists of a skull and jaw with most of the teeth intact. The second species, ''A. robustus'', is known from a skull also from the Pisco Formation in the Cerro la Bruja locality, named MUSM 2182, dating back to the
Serravallian The Serravallian is, in the geologic timescale, an age or a stage in the middle Miocene Epoch/Series, which spans the time between 13.82 Ma and 11.63 Ma (million years ago). The Serravallian follows the Langhian and is followed by the Tortonian ...
–Tortonian stages of the Miocene, older than at least 9.2 mya. A second ''A. deinodon'' specimen of a right
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is nam ...
from the Aguada de Lomas locality was named MNHM F-PPI 272 and dated to the Messinian 6.9–6.7 mya, though it is possible it actually represents ''A. robustus''. MUSM 1399 of a skull with the front half of the snout missing from the Cerro La Bruja locality was referred to ''Acrophyseter'' but has not been given a species designation. ''A. robustus'' had a more pointed snout, straighter teeth, a decreasing width of the mandible from front to back, a groove on the side of its snout, and a more well-defined supracranial basin which housed the
melon A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a " pepo". Th ...
organ than ''A. deinodon''. The
genus name 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 nomenclat ...
''Acrophyseter'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
–meaning acute, which describes the short, pointed, upturned snout–and –which is the genus name for the modern sperm whale ''Physeter macrocephalus''. 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 ...
''deinodon'' is from the Greek –meaning terrible–and –tooth.


Description

The body length range of ''Acrophyseter'' lies within . ''A. deinodon'' was estimated to be using the distance between the cheek bones in comparison to the dimensions of ''Zygophyseter'', which is relatively small, being the smallest of the raptorial sperm whales. Unlike the modern sperm whales, ''A. deinodon'' had teeth on both its upper and lower jaws. The teeth were robust and deeply set into the
roots A root is the part of a plant, generally underground, that anchors the plant body, and absorbs and stores water and nutrients. Root or roots may also refer to: Art, entertainment, and media * ''The Root'' (magazine), an online magazine focusing ...
, particularly the front teeth, the tooth roots were comparatively thick with the thin
tooth crown In dentistry, crown refers to the anatomical area of teeth, usually covered by enamel. The crown is usually visible in the mouth after developing below the gingiva The gums or gingiva (plural: ''gingivae'') consist of the mucosal tiss ...
. The front teeth were more conical than the back teeth. The lower back teeth were close together, and the space between the teeth increased from front to back, suggesting they were used for shearing, unlike the suction-feeding modern-day sperm whales which lack teeth in their upper jaws. The front teeth were more worn on the sides, whereas the bottom teeth were more worn along the middle. It had 12 teeth in the upper jaw and 13 teeth in the bottom jaw, and like other raptorials, it had
tooth enamel Tooth enamel is one of the four major tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the crown. The other major tissues are dentin, ...
. 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 ...
e bore three teeth, and the
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. T ...
e had nine teeth. The last bottom teeth may have contacted the roof of the mouth. Discovered along the teeth sockets was
buccal exostoses A buccal exostosis is an exostosis (bone prominence) on the buccal surface (cheek side) of the alveolar ridge of the maxilla or mandible. More commonly seen in the maxilla than the mandible, buccal exostoses are considered to be site specific. Exi ...
, bony growths, which may have developed during biting to strengthen the teeth, acting as buttresses. The back teeth had larger buccal exostoses as they experienced more pressure during biting. The tooth count of ''A. robustus'' is unknown though thought to be similar or the same.
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, ...
was continually added to the teeth as they were growing, as in
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 ...
s (''Orcinus orca''). Like other sperm whales, ''Acrophyseter'' had a deep basin on the top of its skull, the supracranial basin. This basin in ''Acrophyseter'' overhung the
orbit In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as ...
around the eye, but did not extend onto the snout, unlike in other raptorials. Unlike later species of sperm whales, ''Acrophyseter'' had two nostrils. 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 were as high as they were long, unlike in ''Zygophyseter'' and ''Brygmophyseter'', which displaced the
brow ridge The brow ridge, or supraorbital ridge known as superciliary arch in medicine, is a bony ridge located above the eye sockets of all primates. In humans, the eyebrows are located on their lower margin. Structure The brow ridge is a nodule or crest ...
. The area between 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 ...
, which connects the jaw with the skull, and the teeth was probably where the
masseter muscle In human anatomy, the masseter is one of the muscles of mastication. Found only in mammals, it is particularly powerful in herbivores to facilitate chewing of plant matter. The most obvious muscle of mastication is the masseter muscle, since it ...
s were. The brow ridge slopes down at an angle of around 55°. The
nuchal crest The nuchal lines are four curved lines on the external surface of the occipital bone: * The upper, often faintly marked, is named the highest nuchal line, but is sometimes referred to as the Mempin line or linea suprema, and it attaches to the epi ...
on the back side of the skull had overhung the supracranial basin. The
cheekbones In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygoma ...
were thin plates which limited the ear canals. The snout was short and, unlike in other sperm whales, had a distinct upward curve. Unlike other sperm whales, the top of the premaxillae near 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 max ...
lacked a deep groove. The left nostril was five times bigger than the right nostril, measuring across respectively.


Paleoecology

The short and pointed snout, coupled with the robust, curved front teeth suggests ''Acrophyseter'' targeted large prey and perhaps used their back teeth for shearing. The Sud-Sacaco locality of the Pisco Formation has yielded several marine vertebrates, which the ''Acrophyseter'' may have preyed upon: the whale '' Piscolithax'', the whale '' Piscobalaena'', the seal '' Acrophoca'', the penguin ''
Spheniscus urbinai The banded penguins are penguins that belong to the genus ''Spheniscus''. There are four living species, all with similar banded plumage patterns. They are sometimes also known as "jack-ass penguins" due to their loud locator calls sounding simil ...
'', the marine sloth '' Thalassocnus natans'', the crocodile '' Piscogavialis'',
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 membe ...
, and the broad-toothed mako shark (''Cosmopolitodus hastalis''). The Cerro La Bruja locality has borne the dolphin '' Brachydelphis'', the dolphin '' Atocetus iquensis'', the kentriodontid dolphin '' Belonodelphis'', an unspecified
beaked whale Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least known groups of mammals because of their deep-sea habitat and apparent low abundance. Only three or four of the 24 species are reasonably well-k ...
, unspecified baleen whales, an unspecified
monk seal Monk seals are earless seals of the tribe Monachini. They are the only earless seals found in tropical climates. The two genera of monk seals, ''Monachus'' and ''Neomonachus'', comprise three species: the Mediterranean monk seal, ''Monachus mona ...
, the penguin ''Spheniscus muizoni'', megalodon, the broad-toothed mako shark, and a species of '' Carcharhinus shark.


Taxonomy

''Acrophyseter'', together with ''
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 ...
'', ''
Livyatan ''Livyatan'' is an extinct genus of macroraptorial sperm whale containing one known species: ''L. melvillei''. The genus name was inspired by the biblical sea monster Leviathan, and the species name by Herman Melville, the author of the famous ...
'', 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 ...
'', belong to a group of
macroraptorial sperm whale Macroraptorial sperm whales were highly predatory whales of the sperm whale superfamily (Physeteroidea) of the Miocene epoch that hunted large marine mammals, including other whales, using their large teeth. They consist of five genera: ''Acrophy ...
s, which have adaptations to hunting large prey. They all have large, deeply rooted teeth coated in enamel in both the upper and lower jaws, unlike the modern sperm whale (''Physeter macrocephalus'') which lacks enamel and teeth in the upper jaw. Raptorials are thought to have either evolved these adaptations from a
basilosaurid Basilosauridae is a family of extinct cetaceans. They lived during the middle to the early late Eocene and are known from all continents, including Antarctica. They were probably the first fully aquatic cetaceans.Buono M, Fordyce R.E., Marx F.G. ...
-like ancestor or independently once or twice within the group. The extinct subfamily Hoplocetinae has been proposed to house this group, alongside the genera ''
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 ...
'', '' 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 ...
'', and '' Hoplocetus''. This subfamily is paraphyletic, in that it does not consist of a common ancestor and all of its descendants.


References

{{Taxonbar, from=Q779770 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 2008