MARINE MAMMAL SCIENCE, 14(3):544-55 1 (July 1998) 0 1998 by the Society for Marine Mammalogy

GENERIC NAMES OF NORTHERN AND SOUTHERN FUR SEALS (MAMMALIA:

OTARIIDAE) ALFRED L. GARDNER

Biological Survey, Biological Resources Division, USGS, National Museum of Natural History,

Washington, D.C. 20560-0111, U.S.A. E-mail: gardnera@nmnh.si.edu

c. BRIAN RonslNs Division of Mammals, National Museum of Natural History,

Washington, D.C. 20560-0108, U.S.A.

ABSTRACT

We have resolved a nomenclatural problem discovered during research on the northern fur seal that concerns the correct generic name for this taxon and for fur seals of the Southern Hemisphere. The unfortunate practice by some 19th-century authors to use names in their Latinized form but to date them from their first appearance as French common names led to the use of Arctocepbah for southern fur seals when the name correctly applies to the northern fur seal, known today as CaLLorbinus ursinus. However, Arctocepbalus and Callorbinus are antedated by Otoes G. Fischer, 1817, which is the earliest available generic name for the fur seal of the northern Pacific. The earliest available generic name for southern fur seals is Halarctus Gill, 1866. To avoid the confusion that would result from replacing the currently used generic names with those required by strict adherence to the Principle of Priority, we have petitioned the International Commission on Zoological Nomenclature to preserve ArctocepbaLus and Callorbinas for the southern and northern fur seals, respectively.

Key words: Arctocepbalas, Callorbinus, Halarctus, Otoes, taxonomy, nomencla- ture, fur seals, generic names.

Linnaeus (1758) placed all seals in the genus Phoca. PCron (1816), the first to formally subdivide the group, placed the eared seals in a separate genus, Otarza, in which he recognized five species. A year later, G. Cuvier (1817, pp. 166-167) referred to the eared seals as “Les Phoques B oreilles extkrieures. (Otaries. Peron.),” and recognized only Phoca jzlbata Gmelin and Phoca urszna Gmelin. The same year and apparently unaware that PCron had already sep- arated eared seals from Phoca Linnaeus under the name Otarza, G. Fischer

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GARDNER AND ROBBINS: FUR SEALS 545

(1817:445) coined the name Otoes for the eared seals, in which he included “Phoca jubata, ursina, Lin. Gmel.,” citing “Otaries Peron. Les phoques i oreilles. Cuv. Regne an. 1. p. 166.”

In 1824 F. Cuvier described fur seals under the French vernacular Arctoc- 4hale and said the type was Phoca ursina. His illustration (plate 4, numbers la , Ib, and lc) is of a skull of the South African fur seal currently known in the literature as Arctocephalus pusillus (von Schreber, 1775). Geoffroy St.-Hilaire and F. Cuvier (1826; sometimes cited as F. Cuvier 1826) were the first to use the generic name Arctocephalus for the fur seals. They (1826:553) identified the type as “l’ours marin, Phoca ursina, Linn.,” and gave the distribution (p. 554) as “les iles Aleutiennes, et . . . iles Malouines, et . . . Cap.” The synonymy was limited to {Phoca ursina Linnaeus), “Ursus marinus, Steller, Noui comment. petrop., 11, p. 331; Buff., Suppl. 6, pl. 47.” Reference to a non-northern Pacific fur seal (Buffon’s 1782, plate 47, a New Zealand fur seal jide Allen 1880, 1902; = Otaria forsteri Lesson, 1828) and the inclusion of the “Malouines” (Falkland Islands) and “Cap” (Cape of Good Hope) in the known distribution indicate nothing more than the belief that there was only one widespread species of fur seal.

Allen (1870, 1880, 1902, 1905), Gill (1866), Gray (1866a,b, 1869, 1871, 1874), Peters (1 866), and Trouessart (1 897, 1904) dated Arctocephalus from F. Cuvier’s (1824) use of the common name Arctocghale, following the unfortu- nate practice of using names in their Latinized form, but dating them from their first appearance as French vernaculars (see Palmer 1904:39 for discus- sion). As argued earlier by Gray (1866b346, 1869:267), Allen (1880) consid- ered F. Cuvier’s (1824) illustration to apply equally to the description in Geof- froy St.-Hilaire and F. Cuvier (1826) because only Southern Hemisphere fur seals were then available as specimens in European museums. However, Geof- ftoy St.-Hilaire and F. Cuvier (1826) neither referred to F. Cuvier (1824) nor illustrated the species they called Arctocephalus ursinus. Therefore, the objective basis for their name combination is Phoca ursina Linnaeus, 1758, which is based solely on Steller’s (175 1) sea bear from the Aleutian Islands. Arctocephalus dates from 1826, and F. Cuvier’s description and illustration (plate 4) in 1824 have no bearing on the identity of Arctocephalus ursinus (Linnaeus). Further- more, Phoca ursina Linnaeus was clearly stated by Geoffroy St.-Hilaire and F. Cuvier (1826:553) to be the type species of Arctocephalus and, because its sole basis is Steller’s (1751) sea bear “Ursus marinus,” the name combination Arc- tocephalus ursinus (Linnaeus, 1758) can apply only to the northern fur seal.

Gray (1859c:359) proposed Callorhinus as a monotypic genus for the north- ern fur seal with Arctocephalus ursinus Gray, 1859 (=Phoca ursina Linnaeus) as the type species. Next, four genus-group names were proposed in 1866 for southern fur seals in the following sequence: Halarctos Gill, 1866; Arctophoca Peters, 1866 as a subgenus of Otaria; and Euotarza Gray, 1866c and Gypsophoca Gray, 18666, as subgenera of Arctocephalus (as used by Gray 1859c, 1866a,b,c).

Gill (1866), believing Arctocephalus to be the correct generic name for the northern fur seal, recognized Phoca ursina Linnaeus as the type species; how- ever, he dated Arctocephalus from F. Cuvier (1824), not from Geoffroy St.-

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Hilaire and F. Cuvier (1826) when the name technically became available. Gray (18668) quickly disagreed with Gill’s (1866) designation of Phoca tlrsina as the type of Arctocephaltls. Subsequently, Gill (1871, 1872) accepted Gray’s (1859a, 18666) and Allen’s (1870) opinions on the origin and application of Arctorephalzls. Sclater (1900) and others have identified ursina Linnaeus as the type species of Arctocephalus, but they also erred in dating the name from 1824.

Palmer (1 892), believing Callorhinus Gray to be preoccupied by Callirhintls Blanchard, 1850 (a coleopteran) and Callirhinzls Girard, 1857 (a reptile), pro- posed Callotaria as a replacement name. Later, Palmer (1 90 1 : 134) reported on the name Otoes G. Fischer, 1817 and wrote, “Phoca ursina Gmelin (= Phoca ursina Linn.), is the northern fur seal of Bering Sea and, as the only identifiable species in the group, may be considered the type of Otoes.” Upon discovering the name Otoes, Palmer (1 90 1) believed his Callotaria was unnecessary as a replacement name because Otoes was available and antedated Callorbinas Gray.

Allen ( 1 9 0 2 ~ 1 6 ; 1905) strongly disagreed with Palmer’s (1901) designa- tion, pointing out that the components of jtlbata as referenced by G. Cuvier (1817) are identifiable and that G. Cuvier’s use of tlrsina was also composite. His main argument (1902:117), however, was “It is thus clear that Otoes is unavailable for the Callotaria group, since if one name can ever be considered as a synonym of another, it is evident that Otoes and Otaria holds such a relation.” With few exceptions (e.g., J. B. Fischer 1829, Lesson 1842, Gray 1843) Otoes G. Fischer, 1817 appears to have been overlooked prior to its use by Palmer (1901). Cabrera (1957) followed Allen (1905) in listing Otoes as a synonym of Otaria Ptron.

Palmer (1904:487-488) acquiesced to Allen’s (1902) belief that Otoes was a synonym of Otaria Pkron and gave the type of Otoes G. Fischer as Phoca jzlbata Gmelin. Despite Palmer’s (1904) acceptance of Allen’s (1902; also see Allen 1905) dubious arguments to the contrary, Palmer’s (1901) designation of Phoca tlrsina Gmelin (=Phoca ursina Linnaeus) as the type species of Otoes G. Fischer is valid and restricted Otoes to the fur seals of the northern Pacific. As the earliest available generic name for the northern fur seal, Otoes antedates Callotaria, Callorhinus, and Arctocephalus.

Otaria Ptron and Otoes G. Fischer are not synonyms, because the former represents sea lions (sometimes called hair seals), the latter was restricted to the northern fur seal, and each has a different type species. For many years, Gray (1866a, 1871, 1874) and other authors have used Otaria exclusively for sea lions. Gill (1866) designated Phoca jtl6ata Schreber, 1776, as type species for Otaria; however, Schreber’s jtlbata was composite, including both northern and southern sea lions. Palmer (1904:486) identified “Otaria leonina PCron (=Phoca jtlbata Forster)” as the type species of Otaria Ptron. Allen (1905:105) also designated the sea lion of Forster (which he equated with Otaria leonina Ptron, and with Otaria byronia {Blainville, 18201) as the type species of Otaria PCron. Scheffer (1958:57) considered Ptron’s (1816) equation of Phoca jubata Schreber with Steller’s sea lion “Leo marinus” (=Eumetopias jtlbattls) to be the valid restriction of the composite Phoca jtlbata Schreber, 1776.

GARDNER AND ROBBINS: FUR SEALS 547

The results of our research confirm that: (1) the two oldest generic names applicable to fur seals are Otoes Fischer, 1817 and Arctocephalus Geoffroy St.- Hilaire and F. Cuvier, 1826; (2) Otoes and Arctocephalus are objective synonyms because they have the same type species (Phoca ursina Linnaeus, 1758); ( 3 ) Otoes is the oldest available name for the northern Pacific fur seal known today as Callorhinus ursinus; and (4) Halarctus Gill, 1866 is the oldest available generic name for the fur seals of the Southern Hemisphere known today as species of Arctocephalus.

For nearly a century, with few exceptions, Otoes has been treated as a syn- onym of Otaria PCron. Arctocephalus has been used almost exclusively for Southern Hemisphere fur seals for nearly 140 yr while Callorhinus has been used for the northern fur seal for the same length of time, except for the brief period when replaced by Callotaria. Halarctus has been treated as a junior synonym of Arctocephalus (sensu Gray 18666; Allen 1870, 1880) almost since it was proposed in 1866.

To correctly apply these names according the Principle of Priority (ICZN 1985, Art. 23) would change the name of the northern Pacific fur seal to Otoes ursinus and the Southern Hemisphere fur seals would become species of Hal- arctus. However, Otoes and Halarctus have not been used for nearly a century, and Arctocephalus and Callorhinus are firmly entrenched in the voluminous scientific and popular literature on fur seals. Because we anticipate great con- fusion resulting from replacing these names with Halarctus and Otoes, we have petitioned the International Commission on Zoological Nomenclature (ICZN) to use its plenary powers to suppress Otoes Fischer, 1817 and to place Arcto- cephalus Geoffroy St.-Hilaire and F. Cuvier, 1826 on the Official List of Generic Names in Zoology with type species Phoca pusilla Schreber, 1875. This action would retain Arctocephalus and Callorhinus as valid names consistent with cur- rent usage. Until a decision is rendered, the correct course of action (ICZN 1985, Art. 80) is to retain Arctocephalus and Callorhinus as the valid names according to current usage. If a positive ruling by the ICZN is forthcoming, the generic synonymies of fur seals would be as follows:

Callorhinus Gray, 1 8 5 9 ~

Northern fur seals

Synonymy: Phoca Linnaeus, 1758:37 (part). Otaria PCron, 1816:37, footnote (part). Otoes G. Fischer, 1817:445 (suppressed under the plenary powers of the Inter-

national Commission on Zoological Nomenclature for the purposes of the Principle of Priority, but not for those of the Principle of Homonymy).

Arctoc@hale F. Cuvier, 1824:205 (part; vernacular, unavailable name). Arctocephalus Geoffroy St.-Hilaire and F. Cuvier, 1826:554 (part). Callorhinus Gray, 1859~359 ; type species Callorhinus ursinus Gray, 1859

(=Phoca ursina Linnaeus, 1758), by monotypy.

548 MARINE MAMMAL SCIENCE, VOL. 14, NO. 3, 1998

Callataria Palmer, 1892:156; proposed as a replacement name on the pre- sumption that Callorhinus Gray was preoccupied by Callirhinus Blanchard, 1850 (a coleopteran) and Callirhinus Girard, 1857 (a reptile).

Arctocephalus Geoffroy St.-Hilaire and F. Cuvier, 1826

Southern fur seals

Synonymy: Phoca: Schreber, 1775: pl. 85 (part, not Phoca Linnaeus, 1758). Otaria: Desmarest, 1817:590 (part, not Otaria Peron, 1816). Arctoc4hale F. Cuvier, 1824:205 (part; vernacular, unavailable name). Arctocephalus Geoffroy St.-Hilaire and F. Cuvier, 1826:5 54; type species Phoca

pusilla Schreber, 177 5 by subsequent designation under plenary powers of the International Commission on Zoological Nomenclature.

Halarctus Gill, 1 866:7; type species Arctocephalus delalandii Gray, 1859 (=Phoca pusilla Schreber, 1775), by original designation.

Arctophoca Peters, 1866:276; type species Otaria philippii Peters, 1866, by monotypy (described as a subgenus of Otaria Peron, 1816).

Euotaria Gray, 1866~236 ; type species Arctocephalus nigrescens Gray, 1859 (=Phoca australis Zimmermann, 1783), by monotypy (described as a sub- genus of Arctocephalus Geoffroy St.-Hilaire and F. Cuvier, 1826).

Gypsophoca Gray, 1866~236 ; type species Arctocephalus cinereus Gray, 1866 (=Otaria forsteri Lesson, 1828), by monotypy (described as a subgenus of Arctocephalus Geoffroy St.-Hilaire and F. Cuvier, 1826). Comments on the name Arctocephalus delalandii-Most authors attribute Arc-

tocephalus delalandii to Gray’s (18596) report on the eared seal of the Cape of Good Hope in which the name Otaria delalandii appears in the title. Gray describes this seal under the name Arctocephalus delalandii and in the synonymy listed “Otaria delalandii, F. Cuvier, Dict. Sci. Nat. xxxix, 423; Cuvier, Oss Foss. V. 220, t. 18.f.15, skull.” As pointed out by Allen (1880:213, footnote) the first time the citation “Otaria delalandii, F. Cuvier, Dict. Sci. Nat. xxxix, 423” appeared was in J. B. Fischer (1829:232). This citation was uncritically copied by Gray (1850; 18596; 1866aJ) and Peters (1866). There is no men- tion of any seal on page 423 of the Dictionnaire. Subsequent authors (e.g., Gill 1866; Allen 1880) listed Gray (18593) as the author. Our review of the literature reveals that the earliest correct citation is Otaria delalandii Lesson, 1827. Although attributed to “G. Cuv.,” Lesson is the author of the name; preceding the description he stated (p. 206) “Cette espke, rapportee du Cap par M. Delalande, . . . .”

ACKNOWLEDGMENTS

We are grateful to R. Fisher, J. Mead, C . Sabrosky, and an anonymous reviewer for valuable suggestions and advice during the research and writing of this report.

GARDNER AND ROBBINS: FUR SEALS 549

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Received: 17 October 1995 Accepted: 6 August 1997