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Temporal range: 11.6–7.2 mya Tortonian
Side-view of the skull with the snout facing left, with a flat top and a small upward slope on the right side above the eye socket. All of the teeth are noticeably lighter below a certain point
Cast of a skull at the Museo storia naturale di Pisa in Italy
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Infraorder: Cetacea
Superfamily: Physeteroidea
Family: incertae sedis
Genus: Zygophyseter
Species: Z. varolai
Binomial name
Zygophyseter varolai
Bianucci & Landini, 2006

Zygophyseter varolai is an extinct cetacean that lived during the Tortonian Age of the Late Miocene 11.2 to 7.6 million years ago, and was similar to the modern day sperm whale (Physeter macrocephalus). It is known from a single specimen from the Pietra Leccese Formation in Italy. It was a member of a stem group of fossil macroraptorial sperm whales (often shortened to "raptorial") Brygmophyseter, Acrophyseter, and Livyatan. It grew to be around 6.5 to 7 meters (21 to 23 ft) in length and shared some characteristics with other raptorials, such as large teeth with tooth enamel that were functional in both the upper and lower jaws. It also had a beak, the ability to echolocate prey, and could have probably swum faster than the modern day sperm whale which goes 4 kilometers per hour (2.5 mph). These were probably used in the capture of large prey, such as large fish, seals, and whales. In fact, its common name, killer sperm whale, refers to its feeding habits that would have had a resemblance to the modern day killer whale (Orcinus orca).

























Relationship of Zygophyseter with other sperm whales [1]

The type and only specimen, labelled MAUL 229/1, is of an almost complete skeleton discovered in Southern Italy by geologist Angelo Varola in the marine lime mudstone of the Pietra Leccese Formation near the city of Lecce. It was described in 2006 by geologists Giovanni Bianucci and Walter Landini from the University of Pisa. The genus name Zygophyseter comes from the Latin word zygomaticus, which emphasizes the elongation of the zygomatic process of the only known species Z. varolai, and the term physeter refers to the modern day sperm whale (Physeter macrocephalus) of the family Physeteridae. The species name honors the discoverer.[2][3]

Zygophyseter is part of a fossil stem group of hyper-predatory macroraptorial sperm whales (often shortened to "raptorial") which also includes Brygmophyseter, Acrophyseter, and Livyatan. This group is characterized by having large, functional teeth on both the upper and lower jaw with an enamel coating; whereas the modern sperm whale lacks enamel, teeth in the upper jaws, and functionality in the teeth for catching prey.[1][4] Some fossil remains, mostly teeth, of the genus Scaldicetus were reassigned to these raptorials, including Z. varolai. Scaldicetus is now considered to be a paraphyletic group representing various primitive sperm whale species from different genera. That is to say the genus does not comprise a common ancestor and all of its descendants. It has been proposed that these raptorials be placed into the extinct, paraphyletic subfamily Hoplocetinae, alongside Scaldicetus, Diaphorocetus, Idiorophus, and Hoplocetus.[5] Zygophyseter is more closely related to Brygmophyseter and Acrophyseter than to Livyatan, and the enlarged teeth of this group is thought to have evolved either from a common basilosaurid-like ancestor, or independently once or twice within the group.[6]



A killer sperm whale with a physique similar to a sperm whale, except for a long, black beak. It is hunting tuna
Restoration showing the proposed beak

A characteristic of related raptorials, Zygophyseter had buccal exostoses, bony outgrowths in the alveolar ridge in the mouth, which are thought to have increased their bite force.[2][7][8] Like other raptorials, it had large temporal fossae, probably for supporting strong temporal and masseter muscles, the strongest muscles between the skull and the jaw, meaning this adaptation allowed it to shut its jaws harder. The zygomatic bone (cheekbone) projects anteriorly, indicating it had a beak, which featured an abrupt narrowing; this may have allowed it to clamp down on prey more effectively.[2][6]

The head probably took up 21–23% of the total body size, compared to that of the modern sperm whale which takes up around one-fourth to one-third of the total body size. The blowhole was slanted towards the left side of the animal, and it may have lacked a right nasal passage.[2][9] The falciform process on the squamosal bone was large and ventrally facing; as opposed to the ones in the Kogiidae (Kogia and Praekogia) which are either reduced or absent. These may have been reduced in kogiids due to adaptations to deep sea diving.[10]

Like in modern sperm whales, Zygophyseter had a very large basin above the braincase, known as the supracranial basin, which probably housed the spermaceti organ and the melon. These are used in the generation and focusing of sound for biosonar in the modern sperm whale, indicating Zygophyseter had some mechanisms for biosonar, that is to say this animal could have used echolocation. The zygomatic process of the temporal bone on the cheeks were elongated probably because it contained the spermaceti organ.[11] The skull features a pronounced slope into the supracranial basin.[6] It probably had an echolocation system similar to that of the modern sperm whale, and Zygophyseter may have, in comparison to the echolocative abilities of other modern toothed whales, produced smaller bandwidths and lower center frequencies. This would have made it inept at detecting anything that did not have a diameter of at least 1 meter (3.3 ft), however, unlike the modern sperm whale whose prey is invisible using echolocation, Zygophyseter would have been able to detect prey even though its echolocation system was not that accurate.[2]


The killer sperm whale skull is behind the skull of the other whale, and is seemingly twice as long
Zygophyseter skull (back) next to that of Hemisyntrachelus (front)

Zygophyseter had fourteen teeth in the lower jaw and thirteen in the upper jaw.[9] The curvature of the teeth increased medially, that is, the teeth in the front of the mouth were straighter than the teeth in the back of the mouth. The back teeth featured more wear than the front teeth. Like Brygmophyseter, it had a relatively small crown, making up only 18% of the tooth. Killer whales (Orcinus orca), in comparison, have crowns that make up 20–25% of the tooth. Other characteristics include the presence of the gumline below the crown-root boundary (meaning that part of the root was exposed), and longitudinal grooves on the root. In the type specimen, the teeth ranged in height from 150 to 250 millimeters (5.9 to 9.8 in) with an average height of 175.6 millimeters (6.91 in), and ranged in diameter from 47 to 56 millimeters (1.9 to 2.2 in) with an average of 52.4 millimeters (2.06 in). Like in other raptorials, and unlike in the modern sperm whale, Zygophyseter had tooth enamel.[4][5][12] Like in Acrophyseter, the mandibular foramen takes up about 40% of the lower jawbone. The teeth of the upper jaw form an angle of nearly 120° between the crown and the root, which is possibly a characteristic shared by all raptorials.[6]


Zygophyseter could reach an estimated length of 6.5 to 7 meters (21 to 23 ft), compared to the 12.5-to-18.5-meter (41 to 61 ft) modern sperm whale.[1] It is thought that this whale had twelve thoracic vertebrae and at least ten lumbar vertebra. The type specimen had only 8 thoracic vertebrae preserved, and only the atlas of the cervical vertebrae in the neck. Like in the modern sperm whales, the cervical vertebrae were probably not fused. The centrum of the thoracic vertebrae formed a large and almost pear-shaped central canal which transports nutrients to the spinal cord. The width between the transverse processes (the diagonal projections from a vertebral centrum) of the thoracic vertebrae were 235 millimeters (9.3 in); and the neural spine, the part of the spine that projects away from the centrum, is missing in the type specimen, but it was probably short and thin. The lumbar vertebrae were elongated and may have supported large multifidus and longissimus muscles in the back, likely larger than the modern sperm whale, and so it probably swam faster than the modern sperm whale;[2] the modern sperm whale typically travels horizontally at 4 kilometers per hour (2.5 mph) comparable to other large open-ocean animals.[13] The type specimen had eight caudal vertebrae in the tail.[2]

The animal probably had 12 ribs. The length of the ribs increased from the first to the fifth, then decreased from the fifth to the twelfth; and the width of the ribs decreased from the first to the twelfth, similar to other cetaceans.[2][9]


The common name of Zygophyseter is the "killer sperm whale", due to its similarity to the modern day killer whale (Orcinus orca).[2]

Since the teeth of Zygophyseter are large, exhibit wearing not unlike the teeth of modern day killer whales, and had functionality in both the upper and lower jaws, it was likely a macropredator that fed on large fish, cephalopods, seals, and small whales and dolphins.[5] The position of the condyloid process between the jaw and the skull, like in the modern sperm whale, allowed it to open its jaw wider in order to grab large prey. Its apparent similarity to the feeding habits of the killer whale gave it its nickname "killer sperm whale."[2] The conical front teeth may have functioned as a means for gripping onto prey, and the back teeth were used for cutting. However, neither stomach contents nor cut-marks on the bones of prey species have been discovered, and thus its diet is speculative.[7]

The Z. varolai specimen from the Pietra Leccese Formation dates back to the Tortonian Age of the Late Miocene Epoch, around 11.6 to 7.2 million years ago (mya), and most likely inhabited the Paratethys Sea.[2] This formation has also unearthed the remains of several other large vertebrate species. Ancient sirenians of the genus Metaxytherium were apparently common throughout the ancient Mediterranean sea.[14] Many fish remains of teleost fish, rays, and at least twenty species of sharks have been discovered, such as the tiger shark (Galeocerdo cuvier) and the extinct Carcharocles megalodon.[15] Three species of turtles have been identified: Trachyaspis lardyi, Procolpochelys melii, which are both ancient marine turtles, and Psephophorus polygonus, an ancient leatherback sea turtle.[16] Aside from Z. varolai, two other cetacean species have been described from this formation: the oldest known gray whale Archaeschrichtius ruggieroi, and a species of beaked whale Messapicetus longirostris.[17]

See also[edit]


  1. ^ a b c Berta, A. (2017). The Rise of Marine Mammals: 50 Million Years of Evolution. Baltimore, Maryland: Johns Hopkins University Press. p. 112–113. ISBN 978-1-4214-2326-5. 
  2. ^ a b c d e f g h i j k Bianucci, G.; Landini, W. (2006). "Killer Sperm Whale: a New Basal Physeteroid (Mammalia, Cetacea) from the Late Miocene of Italy". Zoological Journal of the Linnean Society. 148: 103–131. doi:10.1111/j.1096-3642.2006.00228.x. 
  3. ^ Zygophyseter varolai at (retrieved 11 November 2017)
  4. ^ a b Reumer, J. W. F.; Mens, T. H.; Post, K. (2017). "New Finds of Giant Raptorial Sperm Whale Teeth (Cetacea, Physeteroidea) from the Westerschelde Estuary (Province of Zeeland, the Netherlands)" (PDF). Deinsea. 17: 32–38. 
  5. ^ a b c Toscano, A.; Abad, M.; Ruiz, F.; Muñiz, F.; Álvarez, G.; García, E.; Caro, J. A. (2013). "Nuevos Restos de Scaldicetus (Cetacea, Odontoceti, Physeteridae) del Mioceno Superior, Sector Occidental de la Cuenca del Guadalquivir (Sur de España)" [New Remains of Scaldicetus (Cetacea, Odontoceti, Physeteridae) from the Upper Miocene, Western Sector of the Guadalquivir Basin]. Revista Mexicana de Ciencias Geológicas (in Spanish). 30 (2). 
  6. ^ a b c d Lambert, O.; Bianucci, G.; de Muizon, C. (2017). "Macroraptorial Sperm Whales (Cetacea, Odontoceti, Physeteroidea) from the Miocene of Peru". Zoological Journal of the Linnean Society. 179: 404–474. doi:10.1111/zoj.12456. 
  7. ^ a b Lambert, O.; Bianucci, G.; Beaty, B. (2014). "Bony Outgrowths on the Jaws of an Extinct Sperm Whale Support Macroraptorial Feeding in Several Stem Physeteroids". Naturwissenschaften. 101 (6): 517–521. Bibcode:2014NW....101..517L. doi:10.1007/s00114-014-1182-2. 
  8. ^ Marx, F. G.; Lambert, O.; Uhen, M. D. (2016). Cetacean Paleobiology. John Wiley and Sons. pp. 371–372. ISBN 978-1-118-56155-3. 
  9. ^ a b c Lambert, O.; Bianucci, G.; de Muizon, C. (2008). "A New Stem-Sperm Whale (Cetacea, Odontoceti, Physeteroidea) from the Latest Miocene of Peru". Palevol. 7 (6): 361–369. doi:10.1016/j.crpv.2008.06.002. 
  10. ^ Lambert, O. (2008). "Sperm whales from the Miocene of the North Sea: A Re-Appraisal". Sciences de la Terre. 78: 277–316. 
  11. ^ Cranford, T. W.; Amundin, M.; Norris, K. S. (1996). "Functional Morphology and Homology in the Odontocete Nasal Complex: Implications for Sound Generation". Journal of Morphology. 228 (3): 223–285. doi:10.1002/(SICI)1097-4687(199606)228:3<223::AID-JMOR1>3.0.CO;2-3. PMID 8622183. 
  12. ^ Fitzgerald, E. M. G. (2011). "A Fossil Sperm Whale (Cetacea, Physeteroidea) from the Pleistocene of Nauru, Equatorial Southwest Pacific". Journal of Vertebrate Paleontology. 31 (4): 929–931. doi:10.1080/02724634.2011.579670. JSTOR 25835890. 
  13. ^ Whitehead, H. (2003). Sperm Whales: Social Evolution in the Ocean. University of Chicago Press. p. 104–110. ISBN 978-0-226-89517-8. 
  14. ^ Bianucci, G.; Landini, W.; Varola, A. (2003). "New Records of Metaxytherium (Mammalia: Sirenia) from the Late Miocene of Cisterna Quarry (Apulia, Southern Italy)" (PDF). Bollettino della Società Paleontologic a Italiana. 42: 1–2. 
  15. ^ Capasso, L. (2016). "The Fossil Fish of Salento: A History of their Discovery and their Study". Thalassia Salentina. 38: 27–64. doi:10.1285/i15910725v38p27. 
  16. ^ Chesi, F.; Delfino, M.; Varola, A.; Rook, L. (2007). "Fossil sea turtles (Chelonii, Dermochelyidae and Cheloniidae) from the Miocene of Pietra Leccese (late Burdigalian-early Messinian), Southern Italy" (PDF). Geodiversitas. 29 (2): 321–333. 
  17. ^ Bianucci, G.; Collareta, A.; Post, K.; Lambert, O. (2016). "A New Record of Messapicetus from the Pietra Leccese (Late Miocene, Southern Italy): Antitropical Distribution in a Fossil Beaked Whale (Cetacea, Ziphiidae)". Rivista Italiana di Paleontologia e Stratigrafia. 122 (1): 63–74. doi:10.13130/2039-4942/6930. 

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