MARINE MAMMAL SCIENCE, 9(4):372-381 (October 1993)

0 1993 by the Society for Marine Mammalogy

THE SOUTHERN SEA LION,

OTARIA BYRONIA OR OTARIA FLAVESCENS?
DIEGO H. RODRIGUEZ

RICARDO 0. BASTIDA
Consejo National de Investigaciones Cientificas y Tecnicas (CONICET), and
Departamento de Ciencias Marinas, Facultad de Ciencias Exactas y Naturales,
Universidad National de Mar de1 Plats, Dean Funes 3350,
(7600) Mar de1 Plata, Republica Argentina

ABSTRACT

The status of the specific name of the Southern sea lion is controversial;
currently two names are in use: Otaria flavescens (Shaw, 1800) and Otaria
byronia (de Blainville, 1820). The former was based on a two-foot-long yellowish
otariid collected in the Strait of Magellan and the latter on a skull stated to
have been collected on the Tinian Island (Marianas Archipelago); unfortunately
both holotypes are lost. Otaria flavescens has date priority over Otaria byronia,
but its identification was uncertain. This paper focuses on analyzing the main
features of Shaw’s holotype (body length, ear size, and hair color), which fit
with the features of a Southern sea lion pup. As this was a tangible specimen
from a valid locality and has priority over other available epithets, the specific
name of the Southern sea lion should be Otaria flavescens (Shaw, 1800).
Key words: South American otariids, Southern sea lion, nomenclature, Otaria
flavescens (Shaw, 1800), Otaria byronia (de Blainville, 1820), body length, ear
size, hair color.

The specific name of the Southern sea lion has been under discussion for
many years. Currently, two names are in use, Otaria flavescens (Shaw, 1800)
and Otaria byronia (de Blainville, 1820).
Phoca leonina Molina, 1782 was the first description that was clearly of a
Southern sea lion because this description referred to the abundant mane, which
is typical of adult males of the genus Otaria Peron, 1816. Unfortunately the
name Phoca leonina had already been used by Link in 1758 for the Southern
elephant seal. Thus the valid species name would be the next valid epithet
applied to this taxon.
Shaw (1800), based on Pennant (1793), described a small specimen of an
otariid collected in the Strait of Magellan (52”3O’S, 68”3O’W) and stored in
the Leverian Museum. It was named Phoca flavescens by Shaw.
In the same year, 1800, Phoca aurita Bechstein and Phoca flavescens Retzius

372
 RODRIGUEZ AND BASTIDA:SOUTHERN SEA LION 373

Table 1. Main features of Phoca flavescens Shaw.

Feature Pennant 1793 Shaw 1800

Total length “. . rather more than two feet . .” “. . not much exceeding two
feet . . .”
“
Ear length “. an inch long .” . about an inch long . . .”
“
Color . . whitish or cream colour . . .” “. . uniform pale yellow or deep
cream colour without any varie-
gation . .”
Hair length “. . . longish . .” -

Vibrissae “. . very long and whitish . .” “. . . long and whitish , . .”

were published and later related to the synonymy of Otaria. The former was a
German version of Pennant’s work and was rejected in favor of Phoca flavescens
Shaw by Cabrera (1940); the latter name applied, at least in part, to the grey
seal Halichoerus grypus (Osgood 1943). Both names have never subsequently
been used for Otaria and taking into account the generalized use of Otaria
flavescen.s (Shaw, ZSOO), they meet the requirements for considering their sup-
pression (Article 79c of the International Code of Zoological Nomenclature,
Third Edition 1985).
Twenty years later, de Blainville (1820) described his Phoca byronia on the
basis of a skull stated to have been collected on Tinian Island, Marianas Ar-
chipelago (15”N, 146”W), and stored in the Royal College of Surgeons of
London. The skull was destroyed during World War II (Scheffer 1958). The
type locality is obviously incorrect because no otariids have ever been recorded
in this area, but some features, such as the prolonged roof of the palate, are
characteristic of the Southern sea lion skull.
Phoca flavescens Shaw is the earliest available name, with an appropriate type
locality but uncertain identification, whereas the type locality of Phoca byronia
de Blainville is incorrect but its identification is relatively easy (see Ring 1978).
To accept Phoca byronia, the name Phoca flavescens should be previously rejected
(Article 23 of the International Code of Zoological Nomenclature, Third Edition
1985). Consequently, the problem focuses on the identity of Shaw’s holotype,
which unfortunately is lost.

BACKGROUND OF THE CURRENT ISSUE

The relevant features of the original description of Phoca flavescens Shaw are
summarized in Table 1. Four revisions have been made regarding the identity
of this specimen.
Cabrera (1940) states the Southern sea lion is the only otariid in the Magellan
Strait with a yellowish color during youth. Vaz Ferreira (1984) considers that
this color is found in some pups after the first molt and also in prematurely
born specimens of Otaria. Both authors conclude that the holotype corresponds
to a very small Otaria pup.
 374 MARINE MAMMAL SCIENCE,VOL.9,NO.4,1993

Table 2. Neonatal lengths of Southern sea lions (n = 654) and South American fur
seals (n = 34); measurements are range in centimeters. (Sources: Turner 1887, Hamilton
1934, Carrara 1952, Vaz Ferreira 1982a, Vaz Ferreira and Achival 1979, Ponce de Leon
1983, Pinedo 1986, Vaz Ferreira and Ponce de Lebn 1987, Oliva 1988, Contreras et
al. 1988 and Capozzo et al. 1991.)

Range
Species Males Females
Southern sea lion 73-100 62-96
South American fur seal 56-73 57-72

King (1978) regards the holotype as unidentifiable. Oliva (1988) concludes

that Shaw’s Yellow seal was a fur seal pup, probably belonging to the antarctic
fur seal Arctocephalus gazella (Peters, 1875). Both these authors generalize on
the color and size of Otaria pups holding that after the first molt all specimens
are darker and larger than Phoca flavescens Shaw, 1800. The size of the ears
and the length of the hair are also questioned by Oliva (1988), who considers
that these could not correspond to those of the Southern sea lion.

RELATIONSHIP BETWEEN THE TYPE LOCALITY AND OTARIID

BREEDING COLONIES IN THE MAGELLAN STRAIT

The small size of the holotype shows that it could only have been a young
of the otariid species that breed in the proximity of the Magellan Strait. Surveys
during breeding seasons (Carrara 195 2, Vargas and Torres 1976, Sielfeld et al.
1978), and anthropological studies (Schiavini 1987, Vidal and Winograd 1987,
Orquera and Piana 1990) have demonstrated the exclusive presence of the
Southern sea lion and the South American fur seal, Arctocephalus australis
(Zimmermann, 1783), in the southern tip of South America for the last 6,000 yr.
If the holotype had been an antarctic fur seal pup, as suggested by Oliva
(1988), it would require that this species also bred in South America. Apparently
based on a single record of a vagrant juvenile Arctocephalus gazella reported by
Texera (1974) and Payne (1979), Oliva considers these three otariid species to
be sympatric in the Magellan area, even though the breeding colonies of species
of the genus Arctocephalus are usually strictly allopatric (Repenning et al. 197 1,
Rice 1977, Croxall and Gentry 1987). A. gazella is a species whose breeding
is confined to islands close to the Antarctic Polar Front and thus whose nearest
breeding locality (South G eorgia) is 1,500 km away from South America;
furthermore, antarctic fur seal females do not venture farther than 150 km from
their pupping colonies during summer (Croxall and Pilcher 1984, Croxall et
al. 1985, Kooyman et al. 1986). When the pups finally enter the water after
weaning, they are much larger and heavier than the holotype of Phoca flavescens
(Doidge et al. 1984, Doidge and Croxall 1989). In short, the presence of
breeding groups of Arctocephalus gazella in South America is very improbable.
The appearance of vagrant antarctic fur seals outside the species’ normal range
is very recent and probably related with its extraordinary population increase
 RODRiGUEZ AND BASTIDA:SOUTHERN SEA LION 375

Table 3. External ear lengths (mm) of Southern sea lions (n = 11) and South American
fur seals (n = 16); measuremenrs were taken from males and females of all age classes
for both species.

Value Southern sea lion South American fur seal

Mean 25.65 46.40
SD 4.78 4.51
Mean f 1.96 SD 16.28-35.01 37.56-55.24
Observed range 16.00-30.00 40.00-55.00

(Payne 1977, 1978; Laws 1985). Consequently there is no indication suggesting

that the species bred in South America during the pre-exploitation period,
when population levels seem to have been lower than current ones and when
there were no spatial restrictions in the antarctic archipelagos (McCann and
Doidge 1987).

GENERAL FEATURES OF MAGELLANIC OTARIIDS IN

RELATION TO SHAW'S DESCRIPTION

Length at birth--The size of newborn pups is quite variable in both species.

A 2-ft (62-cm) animal could either be a Southern sea lion or a South American
fur seal (Table 2). In the former species the first molt takes place approximately
during the fourth week after birth (Ximenez 197 5, Vaz Ferreira 1982a, Crespo
1988) when the length of the pups has not increased very much. To date, it is
not known what regional differences exist in neonate size in relation to the
shortening of the reproductive season at high latitudes (Campagna 1985).
Since the Phoca flavescens Shaw was a mounted specimen, its skin may have
shrunk. The mean shrinking of Otaria pup pelts is 10 cm, with values ranging
from 6 to 19 cm (Carrara 1952). The length gain during the first month of
life is almost identical to the shrinking of the skin because Southern sea lion
pups grow approximately 3-4 mm daily (Cappozzo et al. 199 1). Consequently
a length of 62 cm would be consistent with the mounted body of a molted
Southern sea lion pup.
Ear size-Among the four authors who analyzed the specific name of Otaria,
only Oliva (1988) indicated that the size of the ear was relevant to the iden-
tification of the holotype. She considered that the stated size was excessive for
a Southern sea lion, although she did not take this characteristic into account
for Arctocephalus australis.
The ear length in both species of Magellanic otariids is variable, but with no
overlap between their values, being significantly smaller in the Otaria we mea-
sured (t = - 11.33, P < 0.00 1; Table 3). The approximate size reported for
the holotype (25.4 mm) lies completely outside the range of Arctocephalus
australis, being smaller even than in 8-mo-old fetuses (32 mm, unpublished
data). On the other hand, although the reported ear length of the holotype
matches rather larger animals than the nominal length of Phoca flauescensShaw,
it corresponds perfectly with the range of Otaria.
376 MARINE MAMMAL SCIENCE, VOL. 9, NO. 4, 1993

Figure 1. Southern sea lion pup found in Península Valdés (Argentina) in October
1988; standard length = 55 cm; color: Munsell (1985) Soil Types 8/6, 7/6 (yellow),
6/6 (brownish yellow) and 5/6 (yellowish brown). Above: general view of the specimen
(horizontal scale = 10 cm). Below: detail of the hair.
 RODRIGUEZ AND BASTIDA: SOUTHERN SEA LION 377

Color-Newborn Southern sea lion pups are practically black. They turn to
dark brown after the first molt, but some specimens show much lighter shades.
Colors of juveniles and adults vary from blackish brown to very light brown
(Hamilton 1934, Aguayo and Maturana 1973, Vaz Ferreira 1982a, King 1983),
while some specimens also vary from pale cream to dull yellow (e.g., Hamilton’s
phases 4 and 5). Although these animals are not pups, it is clear that very
similar colors to that of Shaw’s Yellow Seal are common in the Southern sea
lion. The finding of a 55-cm yellowish pup in Peninsula Valdes (43”15’S,
65”15’W) (Fig. 1) confirms the existence of light-coloured Otaria, as described
by Vaz Ferreira (1984), and fits perfectly with the eighteenth century description
of a 2-ft yellow seal.
Yellowish animals have never been reported for Arctocephalus australis. Only
grey tints similar to that after the first molt, but lighter than the black birthcoat,
were observed by the authors in fetuses of this species. Adult South American
fur seals have dark greyish shades (Vaz Ferreira 19826) but they are never
yellowish as the holotype was. Very similar is the case of Arctocephalus gazella
pups, which are black before and silver-grey after the first molt (J. P. Croxall,
personal communication).
Albinism is rare in Otaria (Lopez and Lopez 1984) and Arctocephalus aus-
tralis (Vaz Ferreira 1984). When it occurs the specimens are completely white,
but different from the white antarctic fur seals which have pinkish brown patches
on the snout, eyes, ears, neck, and flippers (Bonner 1968, Cardenas and Yanez
1983). Shaw’s Yellow Seal is highly unlikely to have been a completely or
partially white animal, given the difference noted by the author between the
whitish vibrisae and the uniform pale yellow body.
Hair length--Oliva (1988) states that the longish hair described by Pennant
(1793) does not correspond to any developmental stage of the Southern sea
lion. However, longish, without any indication of what would be considered
long or short, is almost worthless. Shaw himself, who described as short the
hair of such different species as the common seal, the ribbon seal, the Mediter-
ranean monk seal, the Steller sea lion, and the South American fur seal, clearly
attached no special value to the hair length. Not surprisingly, this characteristic
was ignored by Cabrera (1940), King (1978), and Vaz Ferreira (1984).

OPINIONS

Apart from the four reviews already mentioned, various other opinions about
Shaw’s description have been published since the nineteenth century.
Gray (1868) described the otariid species found in South America and the
Falkland (Malvinas) Islands, where he erroneously believed two species of sea
lions (or hair seals) lived: Otaria jubata and Phocarctos hookeri. As regards the
characteristics of Phoca flavescens Shaw, he stated that “. . . This is very probably
the young of the hair seal of the Falklands . . . which is of a yellowish colour.
Pennant does not mention the want of the under fur . , .” and placed Shaw’s
holotype in the synonymy of Phocarctos hookeri as ‘I. . . young or albino? . . .”
Allen (1880, 1902, 1905) found Phoca flavescens too unsatisfactory and
378 MARINE MAMMAL SCIENCE.VOL. 9. NO. 4. 1993

adopted byronia. Nevertheless he conceded that “. . . the locality, if correctly

indicated, leads to the inference that it was more like a sea lion than a fur seal;
and this being the case it may be hypothetically referred to the genus Otaria,
to which it has been provisionally assigned by the majority of writers during
the last fifty years . . .” and that “. . . from its size, colour and habitat it is
presumably referable to Otaria, but it has been referred by Gray to his Phocarctos
hookeri. . . .” However, he later regarded the genera Phocarctos and Otaria as
restricted to Australasia and South America respectively. This point of view was
followed by Hamilton (1934) and Sivertsen (1956).
After Cabrera’s review, Osgood (1943) agreed that Phoca javescens Shaw
“
. . . was based on a preserved specimen and, although the description is lacking
in details, it contains no contradictory matter. . ” Scheffer (195 8) agreed with
Cabrera, but he preferred the use of byronia; Hershkovitz (in Rice 1977) also
shared Cabrera’s conclusions, stating that Shaw’s holotype “. . . was adequately
described and figured, is perfectly identifiable and has a valid type locality. Its
name has priority, usage and currency. . . .” These arguments were also adopted
by Pine et al. (1978) and Mattlin (1985).

CONCLUSIONS

Shaw’s holotype, collected in the Strait of Magellan, could only have been a
newborn pup of Otaria or Arctocephalus australis; its body size (circa 62 cm)
could correspond to either species, but the uniform yellowish color is found
exclusively in some molted pups of the Southern sea lion. Ear length, though
not matching well with the described body length, lies within the recorded range
of Otaria, but outside that for Arctocephalus austTraliS.
In view of the existence of Southern sea lion pups with the same characteristics
of Phoca flavescens Shaw, and that the holotype was a tangible specimen from
a valid type locality (originally preserved in the Leverian Museum), described
(General Zoology or Systematic Natural History, Volume I, Part II, pages 260-
26 l), illustrated (idem page 73), and with priority over any other available
epithets, the specific name of the Southern sea lion should be Otaria flavescens
(Shaw, 1800).

ACKNOWLEDGMENTS

We wish to express our gratitude to Dr. John P. Croxall (British Antarctic Survey,
U.K.) for commenting on the manuscript and for his continuous help in our pinniped
studies. The Editor and three referees of Marine Mammal Science made helpful suggestions
to improve the submitted version. Many thanks to Gloria, Jose, Eduardo, and Ricardo
Mendez for allowing us to examine the fur seal and sea lion collections at Oceanario
Mundo Marino (San Clemente de1 Tuyu, Argentina) and to make measurements. This
task was successful due to the skill of the seal trainers Jose Nunez, Hector Canatelli, and
Rat2 Florio, and to the patience of the seals Loli, Salvatore, Noel, Poroto, Gina, Tommy,
Lujh, Malvina, Luz, and Lila. Mrs. Cristina Sarasa kindly translated an early draft into
English, and Mrs. Marcela Tobio (INIDEP, Mar de1 Plata) helped us with the photographic
work.
 RODRiGUEZ AND BASTIDA:SOUTHERNSEA LION 379

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Received: October 8, 1992
Accepted: June 3, 1993