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Physeteridae (sperm whale)

Mammalia - Cetacea - Physeteridae

Taxonomy

Physeteridae was named by Gray (1821). It is extant. It was considered monophyletic by Lambert (2008).

It was assigned to Carnivorae by Gray (1821); to Cete by Bonaparte (1850); to Denticete by Scammon and Cope (1869) and Gill (1871); to Denticete by Gill (1873); to Odontoceta by Ameghino (1889); to Odontocete by Hay (1902) and Hay (1930); to Squaloceti by Abel (1914), Abel (1919), Dal Piaz (1929) and Van Deinse (1931); to Odontoceti by Winge (1921) and Slijper (1936); to Cetacea by Pictet (1853), Newton (1891) and Nowak (1991); to Odontoceti by Flower (1867), Flower (1883), Lydekker (1887), Cope (1889), Cope (1890), Flower and Lydekker (1891), Flores (1895), Weber (1904), Trouessart (1904), Case (1904), Abel (1905), True (1908), Jaekel (1911), Turner (1912), Miller (1923), Zittel (1925), Kellogg (1928), Weber (1928), Ellerman and Morrisson-Scott (1951), Scheffer and Rice (1963), Pilleri (1985), Carroll (1988), Vidal (1991), Benton (1993), Luo and Marsh (1996), Sach and Heizmann (2001) and Mead and Brownell (2005); and to Physeteroidea by Gray (1868), Gill (1872), Simpson (1945), Fraser and Purves (1960), Ginsburg and Janvier (1971), Kasuya (1973), Gondar (1974), Barnes et al. (1985), Muizon (1990), Fordyce and Barnes (1994), Fordyce et al. (1995), Hirota and Barnes (1995), McKenna and Bell (1997), Rice (1998), Fordyce and de Muizon (2001), Rice (2002), Kazár (2002), Geisler and Sanders (2003), Gingerich (2005), Hampe (2006), Bianucci and Landini (2007), Foekens (2008), Whitmore and Kaltenbach (2008), Uhen et al. (2008), Lambert (2008), Rice (2009), Geisler et al. (2011), Valerio and Laurito (2012), Ormezzano and Lanzetti (2014), Aguirre-Fernández and Fordyce (2014), Marx et al. (2016), Collareta et al. (2017), Berta (2017), Alfsen et al. (2021) and Godfrey and Lambert (2023).

Sister taxa

Physodontidae

Synonyms

Ziphiola was named by Vanden Broeck (1874) [Vanden Broeck assigns this taxon to Van Beneden. R. Kellogg's card file says this is a nomen nudum.]. Its type is Ziphiola clepsydra.

It was considered a nomen nudum by Kellogg (1969).

It was assigned to Ziphiina by Vanden Broeck (1874); and to Ziphiidae by Abel (1901).
Ziphiola clepsydra was named by Vanden Broeck (1874) [Vanden Broeck assigns this taxon to Van Beneden.]. It is the type species of Ziphiola.

It was considered a nomen nudum by Kellogg (1969).
Ziphioides was named by Probst (1886) [Sepkoski's age data: T Mi-m].

It was considered a nomen dubium by Fordyce and de Muizon (2001).

It was assigned to Ziphiinae by Trouessart (1898) and Trouessart (1904); to Physeteridae by Palmer (1904); to Ziphiidae by Paquier (1894), Kellogg (1928), Simpson (1945), Mead (1975) and Carroll (1988); to Hyperoodontidae by McKenna and Bell (1997); and to Cetacea by Sepkoski (2002).
Paleophoca was named by Van Beneden (1859). Its type is Paleophoca nystii. It was considered monophyletic by Uhen et al. (2008).

It was synonymized subjectively with Pristiphoca by Trouessart (1898) and Trouessart (1904); it was corrected as Palaeophoca by Palmer (1904) and Zittel (1925); it was considered a nomen dubium by Fordyce and de Muizon (2001); it was considered a nomen dubium by Uhen and Pyenson (2007); it was considered a nomen nudum by Koretsky and Ray (2008).

It was assigned to Monachinae by Hendey (1972); to Phocidae by Van Beneden (1859), Palmer (1904), Kellogg (1922), Zittel (1925) and Carroll (1988); to Hoplocetinae by McKenna and Bell (1997); to Carnivora by Sepkoski (2002); and to Physeteridae by Ray (1977), Kazár (2002) and Uhen et al. (2008).
Physotherium was named by Portis (1886).

It was considered a nomen dubium by Fordyce and de Muizon (2001).

It was assigned to Cetacea by Zittel (1893); to Physeterinae by Trouessart (1898) and Trouessart (1904); and to Physeteridae by Palmer (1904) and Pilleri (1980).
Physotherium sotterii was named by Portis (1886) [University of Turin Museum of Paleontlogy ]. Its type specimen is UTMP 23824, a mandible, and it is a 3D body fossil. Its type locality is Astigiana, which is in a Zanclean marine horizon in the Sabbie Gialle Formation of Italy.

It was considered a nomen nudum by Kellogg (1923).
Scaldicetus lodgei was named by Chapman (1917) [no specimen number listed for the holotype in Chapman 1917]. It is a 3D body fossil. Its type locality is Clifton Bank, Muddy Creek (Vaginella band), which is in a Langhian marine marl in the Port Campbell Limestone Formation of Australia.

It was considered a nomen dubium by Hampe (2006).
Orcopsis was named by Van Beneden (1876).

It was synonymized subjectively with Physeterula by Abel (1905); it was considered a nomen dubium by Fordyce and de Muizon (2001).

It was assigned to Delphinidae by Van Beneden (1884), Flores (1895) and Palmer (1904).
Physodontidae was named by Lydekker (1894). It is not extant. Its type is Physodon.

It was considered an invalid subgroup of Physeteridae by Barnes et al. (1985) and McKenna and Bell (1997).

It was assigned to Physeteroidea by Lydekker (1894).
Ziphioides triangulus was named by Probst (1886) [no type specimen listed in Probst 1886]. It is not extant. Its type locality is Baltringen, which is in a Burdigalian marine horizon in Germany.

It was considered a nomen dubium by Fordyce and de Muizon (2001).
Ziphioides obliquus was named by Probst (1886) [no type specimen listed in Probst 1886]. Its type locality is Baltringen, which is in a Burdigalian marine horizon in Germany.

It was considered a nomen dubium by Fordyce and de Muizon (2001).
Orca semseyi was named by Bockh (1899). It is a 3D body fossil. Its type locality is Salgotarjan, which is in a Miocene marine horizon in Hungary.

It was recombined as Orcinus semseyi by Trouessart (1904); it was considered an invalid subgroup of Physeteridae by Heyning and Dahleim (1988) and Hampe (2006).
Catodontidae was named by Cuvier (1836). It is extant.

It was synonymized subjectively with Physeteridae by Barnes et al. (1985) and McKenna and Bell (1997).

It was assigned to Cete by Gray (1846) and Gray (1850); to Denticete by Gray (1864); to Denticete by Gray (1866); and to Physeteroidea by Gray (1868).
Physeterina was named by Gray (1825). It is extant.

It was corrected as Physeterini by Winge (1921); it was synonymized subjectively with Physeteridae by McKenna and Bell (1997).

It was assigned to Balaenidae by Gray (1825); and to Physeteridae by Bonaparte (1850) and Winge (1921).

Synonymy list

Year	Name and author
1821	Physeteridae Gray p. 310
1825	Physeterina Gray p. 340
1836	Catodontidae Cuvier p. 564
1846	Catodontidae Gray p. 21
1850	Physeteridae Bonaparte p. 1
1850	Physeterina Bonaparte p. 1
1850	Catodontidae Gray p. 44
1853	Physeteridae Pictet p. 386
1859	Paleophoca Van Beneden
1864	Catodontidae Gray p. 231
1866	Catodontidae Gray p. 195
1867	Physeteridae Flower p. 113
1868	Catodontidae Gray p. 3
1868	Physeteridae Gray p. 3
1869	Physeteridae Scammon and Cope p. 20
1871	Physeteridae Gill p. 123
1872	Physeteridae Gill p. 96
1873	Physeteridae Gill p. 27
1874	Ziphiola Vanden Broeck p. 146
1874	Ziphiola clepsydra Vanden Broeck p. 146
1876	Orcopsis Van Beneden p. 489
1883	Physeteridae Flower p. 184
1884	Orcopsis Van Beneden p. 9
1886	Physotherium Portis p. 325 figs. Plate VII
1886	Physotherium sotterii Portis pp. 325-326 figs. Plate VII
1886	Ziphioides Probst
1886	Ziphioides triangulus Probst p. 110 figs. Fig. 7b
1886	Ziphioides obliquus Probst p. 112 figs. Fig. 8b
1887	Physeteridae Lydekker p. 53
1889	Physeteridae Ameghino p. 887
1889	Physeteridae Cope p. 876
1890	Physeteridae Cope p. 606
1891	Physeteridae Flower and Lydekker p. 89
1891	Physeteridae Newton p. 65
1893	Physotherium Zittel p. 502
1894	Physodontidae Lydekker p. 4
1894	Ziphioides Paquier p. 47
1895	Orcopsis Flores p. 12
1895	Physeteridae Flores p. 13
1898	Physotherium Trouessart p. 1054
1898	Ziphioides Trouessart p. 1059
1898	Ziphioides obliquus Trouessart p. 1059
1898	Ziphioides triangulus Trouessart p. 1059
1899	Orca semseyi Bockh p. 43 figs. Figure 1
1901	Ziphiola Abel p. 41
1902	Physeteridae Hay p. 595
1904	Physeteridae Case p. 30
1904	Orcopsis Palmer p. 478
1904	Palaeophoca Palmer p. 505
1904	Physotherium Palmer p. 537
1904	Ziphioides Palmer p. 716
1904	Orcinus semseyi Trouessart p. 771
1904	Physeteridae Trouessart p. 772
1904	Physotherium Trouessart p. 773
1904	Physotherium solterii Trouessart p. 773
1904	Ziphioides Trouessart p. 775
1904	Ziphioides obliquus Trouessart p. 775
1904	Ziphioides triangulus Trouessart p. 775
1904	Physeteridae Weber p. 578
1905	Physeteridae Abel p. 51
1908	Physeteridae True p. 391
1911	Physeteridae Jaekel p. 231
1912	Physeteridae Turner p. 71
1914	Physeteridae Abel p. 221
1917	Scaldicetus lodgei Chapman p. 34 figs. Plate IV, Fig. 6
1919	Physeteridae Abel p. 763
1921	Physeteridae Winge p. 8
1921	Physeterini Winge p. 8
1922	Paleophoca Kellogg p. 111
1923	Physeteridae Miller p. 40
1925	Palaeophoca Zittel p. 78
1925	Physeteridae Zittel p. 86
1928	Ziphioides Kellogg p. 33 figs. Table 1
1928	Physeteridae Kellogg p. 34 figs. Table 1
1928	Physeteridae Weber p. 389
1929	Physeteridae Dal Piaz p. 78
1930	Physeteridae Hay p. 596
1931	Physeteridae Van Deinse p. 33
1936	Physeteridae Slijper p. 549
1945	Ziphioides Simpson p. 101
1945	Physeteridae Simpson p. 102
1951	Physeteridae Ellerman and Morrisson-Scott p. 720
1960	Physeteridae Fraser and Purves p. 112 figs. Figure 26
1963	Physeteridae Scheffer and Rice p. 8
1971	Physeteridae Ginsburg and Janvier p. 172
1972	Palaeophoca Hendey p. 100
1973	Physeteridae Kasuya p. 61
1974	Physeteridae Gondar
1975	Ziphioides Mead p. 750 figs. Table 1
1977	Paleophoca Ray p. 395
1980	Physotherium Pilleri p. 51
1985	Physeteridae Barnes et al. p. 26
1985	Physeteridae Pilleri p. 30
1988	Palaeophoca Carroll
1988	Physeteridae Carroll
1988	Ziphioides Carroll
1990	Physeteridae Muizon p. 297
1991	Physeteridae Nowak
1991	Physeteridae Vidal p. 5
1993	Physeteridae Benton p. 761
1994	Physeteridae Fordyce and Barnes p. 428 figs. Table 1
1995	Physeteridae Fordyce et al. p. 379
1995	Physeteridae Hirota and Barnes p. 455
1996	Physeteridae Luo and Marsh p. 331
1997	Physeteridae McKenna and Bell p. 379
1997	Paleophoca McKenna and Bell p. 380
1997	Ziphioides McKenna and Bell p. 382
1998	Physeteridae Rice
2001	Physeteridae Fordyce and de Muizon p. 179
2001	Physeteridae Sach and Heizmann p. 42
2002	Physeteridae Kazár p. 154
2002	Paleophoca Kazár p. 163
2002	Scaldicetus lodgei Kazár p. 163
2002	Physeteridae Rice p. 231 figs. Table 1
2002	Palaeophoca Sepkoski
2002	Ziphioides Sepkoski
2003	Physeteridae Geisler and Sanders p. 28
2004	Scaldicetus lodgei Fitzgerald p. 206
2005	Physeteridae Gingerich p. 237 figs. Table 15.1
2005	Physeteridae Mead and Brownell p. 737
2006	Physeteridae Hampe p. 65
2007	Physeteridae Bianucci and Landini p. 45 figs. Table 2.1
2008	Physeteridae Foekens p. 7 figs. Table 1
2008	Physeteridae Lambert p. 293
2008	Physeteridae Uhen et al. p. 574
2008	Paleophoca Uhen et al. p. 577
2008	Physeteridae Whitmore and Kaltenbach p. 236
2009	Physeteridae Rice p. 235 figs. Table 1
2011	Physeteridae Geisler et al. p. 5 figs. Table 1
2012	Physeteridae Valerio and Laurito p. 153
2014	Physeteridae Aguirre-Fernández and Fordyce p. 195
2014	Physeteridae Ormezzano and Lanzetti p. 20
2016	Physeteridae Marx et al. p. 123
2017	Physeteridae Berta p. 161
2017	Physeteridae Collareta et al. p. 269 figs. Figure 6
2021	Physeteridae Alfsen et al. p. 809
2023	Physeteridae Godfrey and Lambert p. 63

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
subclass	Synapsida
—	Therapsida	()
infraorder	Cynodontia	()
—	Mammaliamorpha
—	Mammaliaformes
class	Mammalia

Rank	Name	Author
—	Theriamorpha	(Rowe 1993)
—	Theriiformes	()
—	Trechnotheria
—	Cladotheria
—	Zatheria
subclass	Tribosphenida	()
subclass	Theria
—	Eutheria	()
—	Placentalia
—	Boreoeutheria
—	Laurasiatheria
—	Scrotifera
—	Euungulata
—	Artiodactylamorpha
—	Artiodactyla	()
—	Whippomorpha
order	Cetacea
—	Pelagiceti
—	Neoceti
suborder	Odontoceti
superfamily	Physeteroidea
family	Physeteridae
family	Physeteridae

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Fm. Physeteridae Gray 1821 [sperm whale]

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G. †Aulophyseter Kellogg 1927

†Aulophyseter mediatlanticus Cope 1895

†Aulophyseter morricei Kellogg 1927

G. †Diaphorocetus Ameghino 1894

†Diaphorocetus ortegai Lambert et al. 2023

†Diaphorocetus poucheti Moreno 1892

Invalid names: Hypocetus Lydekker 1894 [objective synonym], Mesocetus Moreno 1892 [synonym], Paracetus Lydekker 1894 [objective synonym]

Subfm. †Hoplocetinae Cabrera 1926

Invalid names: Balaenodon physaloides Owen 1846 [nomen dubium], Physetodon baileyi McCoy 1879 [nomen dubium], Physodon leccense Gervais 1872 [nomen dubium]

G. †Idiophyseter Kellogg 1925

†Idiophyseter merriami Kellogg 1925

G. †Idiorophus Kellogg 1925

†Idiorophus bolzanensis Dal Piaz 1916

†Idiorophus patagonicus Lydekker 1894

Invalid names: Apenophyseter Cabrera 1926 [synonym]

G. †Kogiopsis Kellogg 1929

†Kogiopsis floridana Kellogg 1929

G. †Orycterocetus Leidy 1853

†Orycterocetus cornutidens Leidy 1856

†Orycterocetus crocodilinus Cope 1867

Invalid names: Gargantuodon ligerensis Ginsburg 1969 [synonym]

†Orycterocetus quadratidens Leidy 1853

Invalid names: Gargantuodon Ginsburg 1969 [synonym]

G. Physeter Linnaeus 1758

†Physeter antiquus Gervais 1849

Physeter macrocephalus Linnaeus 1758 [sperm whale]

Invalid names: Megistosaurus ensiformis Godman 1827 [synonym], Physeter catodon Linnaeus 1758 [synonym]

†Physeter vetus Leidy 1869

Invalid names: Physeter antiquus Leidy 1853 [objective synonym]

Invalid names: Catodon Linnaeus 1761 [synonym], Cetus Billberg 1828 [synonym], Meganeuron Gray 1866 [synonym], Megistosaurus Godman 1827 [synonym], Orthodon Rafinesque 1815 [synonym], Physalus Lacépède 1804 [synonym]

G. †Physeterula Van Beneden 1877

†Physeterula dubusi Van Beneden 1877

Invalid names: Delphinus acutidens von Meyer 1859 [synonym]

†Physeterula neolassicus Veatch and Stephenson 1911

G. †Placoziphius Van Beneden 1869

†Placoziphius duboisii Van Beneden 1869

G. †Thalassocetus Abel 1905

†Thalassocetus antwerpiensis Abel 1905

Invalid names: Aulophyseterinae Kazár 2002 [empty], Catodontidae Cuvier 1836 [synonym], Orca semseyi Bockh 1899 [invalid subgroup], Orcopsis Van Beneden 1876 [nomen dubium], Paleophoca Van Beneden 1859 [nomen nudum], Physeterina Gray 1825 [synonym], Physeterinae Flower 1867 [empty], Physodontidae Lydekker 1894 [invalid subgroup], Physotherium Portis 1886 [nomen dubium], Scaldicetus lodgei Chapman 1917 [nomen dubium], Ziphioides Probst 1886 [nomen dubium], Ziphiola Vanden Broeck 1874 [nomen nudum]

Diagnosis

Reference	Diagnosis
K. Hirota and L. G. Barnes 1995	A family of Odontoceti having cranium with large supracranial basin, bounded posteriorly by a high, transverse, semicircular nuchal crest and laterally by a maxillary crest; lacking a median crest or a fossa in the supracranial basin; lacrimal and jugal fused; jugular part of zygomatic process short and wide, not reaching posteriorly to contact zygomatic process of squamosal; palatines not forming part of anterior wall of narial passage; narial passages of disparate sizes with right naris being smaller; pterygoid enlarged, without reduplication, hamulus small; zygomatic process of squamosal not flaring laterally from braincase; posterior part of rostrum wide and flat; air-sinus system simple, without preorbital and postorbital lobes; left premaxillary foramen lost; mandibular symphysis elongate and narrow (mandible Y-shaped); teeth single-rooted and with conical crowns, homodont; tympanic bulla relatively small; mastoid process large; accessory ossicle on periotic; all cervical vertebrae except atlas ankylosed; forelimb short, humerus approximately equal in length to radius and ulna; olecra- non process of ulna large and triangular in shape.
M. D. Uhen et al. 2008	The near-cosmopolitan living sperm whale Physeter catodon Linnaeus, 1758, or Physeter macrocephalus Linnaeus, 1758 (see Rice 1998 on nomenclature), is sufficiently disparate from the related living species of Kogia (Kogiidae) to be placed in its own family Physeteridae, or crown-Physeteridae. Many fossil species have been allied to Physeter, but nomenclature is confusing especially above genus-level; crown-stem terminology has yet to be applied to the Physeter lineage. The fragmentary and enigmatic late Oligocene Ferecetotherium kelloggi Mchedlidze, 1970, from Azerbaidjan is the oldest known sperm whale (Barnes, 1985c), making sperm whales the oldest surviving family-level clade of Odontoceti. The distinctive skull form of Physeter macrocephalus reflects the unique morphology of cranial soft tissues, as documented by Raven and Gregory (1933), Fraser and Purves (1960), Heyning (1989a), Cranford (1999, Cranford et al. 1996), and others. Diagnostic bony features for Physeter (namely, for crown-Physeteridae) include: large absolute size; long flattened triangular rostrum with generally vestigial teeth; large supracranial basin with elevated maxillary walls that lie medial to antorbital notches and large slit-like dorsal infraorbital foramina; no facial sagittal crest or elevated postnarial vertex within the supracranial basin; parabolic posterior part of maxillary crest is confluent with nuchal crest of supraoccipital; teeth homodont; periotic with enlarged accessory ossicle; and cervical vertebrae 2-7 fused. Many fossils show some of these features, or others seen in Physeter but not detailed above, and there is little doubt that such fossils are in the Physeter lineage. For example, the Early Miocene Diaphorocetus poucheti (South Atlantic) has a supracranial basin but a more plesiomorphic brain case, which links it firmly with other odontocetes. Many fossil species, for example, in the genus Scaldicetus, retained the plesiomorphic condition of large functional teeth in both the upper and lower jaws (for example Hirota and Barnes, 1995), while others have small or vestigial upper teeth (for example Aulophyseter and Orycterocetus), as does living P. macrocephalus. The more archaic fossil sperm whales had enamel-covered tooth crowns, but in various later Tertiary fossil sperm whales the crowns of the teeth lack enamel, as does P. macrocephalus. The homodonty and enlarged supracranial basin in fossil Physeteridae have led, by analogy with the living P. macrocephalus, to the idea that sperm whales have mainly been deep-diving squid-eaters (e.g. Cozzuol, 1996). Miocene and Pliocene fossil sperm whales include many named species worldwide. Many taxa have been based on isolated teeth of uncertain taxonomic value (e.g. Kellogg 1925a), thus creating problems in understanding the taxonomy and history of the family. Subfamily groupings used within the Physeteridae include Physeterinae, Scaldicetinae, Hoplocetinae and Aulophyseterinae (e.g. Simpson 1945, Kazar 2002). The living P. macrocephalus is not known from the fossil record of North America. Despite reports cited by Dorr and Eschman (1970) and Holman (1995) of Physeter from Great Lakes deposits in Michigan, these specimens have been determined to be of historical (less than 1000 years) age and unlikely to have been naturally deposited (Harrington, 1988).