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Encyclopedia of Marine Mammals, Second Edition 878 Pinnipedia, Overview northern elephant seals. Am. J. Physiol. Regul. Integr. Comp. Physiol. Pinnipeds comprise slightly more than one-fourth (26%) of the 293 , R2376 – R2381 . species diversity of marine mammals (approximately 128 species Iverson , S. J. , Hamosh , M. , and Bowen , W. D. ( 1995 ). Lipoprotein lipase currently recognized). Thirty-three living species of pinnipeds are activity and its relationship to high milk fat transfer during lactation distributed throughout the world: 18 phocids, 14 otariids, and the in grey seals . J. Comp. Physiol. B 165 , 384 – 395 . walrus. One additional species of phocid (Caribbean monk seal) and Keith , E. O. , and Ortiz , C. L. ( 1989 ). Glucose kinetics in neonatal ele- one subspecies of otariid (Japanese sea lion) are reported extinct in phant seals during postweaning aphagia . Mar. Mamm. Sci. 5 , 99 – 115 . Lang , S. L. C. , Iverson , S. J. , and Bowen , W. D. ( 2005 ). Individual varia- historical time. tion in milk composition over lactation in harbour seals (Phoca vitu- lina ) and the potential consequences of intermittent attendance. Can. I. Systematics and Distribution J. Zool. 83 , 1525 – 1531 . Mellish , J. E. , Iverson , S. J. , Bowen , W. D. , and Hammill , M. O. ( 1999 ). A. Otariidae: Fur Seals and Sea Lions Fat transfer and energetics during lactation in the hooded seal: the Of the two groups of seals, the otariids are characterized by the pres- roles of tissue lipoprotein lipase in milk fat secretion and pup blubber ence of external ear fl aps or pinnae, and for this reason they are often deposition . J. Comp. Physiol. B 169 , 377 – 390 . called “ eared ” seals ( Fig. 1 ). Otariids (and the walrus) can turn their Nordoy , E. S. , Stijfhoorn , D. E. , Raheim , A. , and Blix , A. S. ( 1992 ). hindfl ippers forward and use them to walk (discussed further under Water fl ux and early signs of entrance into phase-iii of fasting in gray Adaptations). The Otariidae traditionally are divided into two sub- seal pups . Acta Physiol. Scand. 144 , 477 – 482 . Nord ø y , E. S. , and Blix , A. S. ( 1991 ). Glucose and ketone body turnover families Otariinae (sea lions) and Arctocephalinae (fur seals) although in fasting grey seal pups . Acta Physiol. Scand. 141 , 563 – 571 . recent molecular studies have revealed that neither fur seals nor sea Ortiz , R. M. , Wade , C. E. , and Ortiz , C. L. ( 2001 ). Effects of prolonged lions are monophyletic. Five living genera and species of sea lions are fasting on plasma cortisol and TH in postweaned northern ele- recognized, occurring in both the northern and southern hemispheres: phant seal pups . Am. J. Physiol. Regul. Integr. Comp. Physiol. 280 , Eumetopias jubata (northern or Steller’s sea lion), Neophoca cinerea R790 – R795 . (Australian sea lion), Otaria byronia (Southern sea lion), Zalophus cali- Ortiz , R. M. , Wade , C. E. , and Ortiz , C. L. ( 2003 ). Body water handling fornianus ( Z. c. californianus California sea lion, Z. c. japonicus Japanese in response to hypertonic-saline induced diuresis in fasting north- sea lion, and Z. c. wollebacki Galapagos sea lion), and Phocarctos ern elephant seal pups (Mirounga angustirostris ) . Comp. Biochem. hookeri (New Zealand sea lion; Fig. 2 ). The fur seals, named for their Physiol. A 134 , 423 – 428 . thick, dense fur, include two genera, the monotypic Northern fur seal Ortiz , R. M. , Crocker , D. E. , Houser , D. S. , and Webb , P. M. ( 2006). Angiotensin II and aldosterone increase with fasting in breeding adult (Callorhinus ursinus ) and the southern fur seals (Arctocephalus ), con- male northern elephant seals ( Mirounga angustirostris ) . Physiol. sisting of eight species: A. australis (South American fur seal), A. forsteri Biochem. Zool. 79 ( 6 ) , 1106 – 1112 . (New Zealand fur seal), A. gazella (Antarctic fur seal), A. galapagoensis Pernia , S. D. , Costa , D. P. , and Ortiz , C. L. ( 1989 ). Glomerular fi ltration (Galapagos fur seal), A. philippii (Juan Fernandez fur seal), A. pusillus rate in weaned elephant seal pups during natural, long term fasts. (A. p. pusillus South African fur seal and A. p. doriferus Australian fur Can. J. Zool. (J. Can. Zool.) 67 , 1752 – 1756 . seal), A. townsendi (Guadalupe fur seal), and A. tropicalis (Subantarctic Sharp , J. A. , Cane , K. N. , Lefevre , C. , Arnould , J. P. Y. , and Nicholas , K. fur seal). All of the fur seals except the northern and Guadalupe fur seals R. ( 2006 ). Fur seal adaptations to lactation: Insights into mammary are found in the southern hemisphere. The northern fur seal is found gland function . Curr. Top. Dev. Biol. 72 , 276–308 . in subarctic waters of the North Pacifi c, with the exception of a small Sharp , J. A. , Lefevre , C. , Brennan , A. J. , and Nicholas , K. R. ( 2007). population on San Miguel Island off California ( Fig. 3 ). The fur seal — a model lactation phenotype to explore molecular fac- tors involved in the initiation of apoptosis at involution . J. Mammary Otariid monophyly is well supported based on a both morphologic Gland Biol. Neoplasia 12 , 4 7 – 5 8 . as well as molecular data. Morphologic and molecular studies con- Tedman , R. A. ( 1983 ). Ultrastructural morphology of the mammary- sistently position Callorhinus as the earliest diverging extant otariid gland with observations on the size distribution of fat droplets in milk ( Fig. 4 ). Extinct otariids Pithanotaria , Thalassoleon , and Hydractos of the Weddell seal Leptonychotes weddelli (Pinnipedia) . J. Zool. are sequential sister taxa to Callorhinus . Molecular results strongly 200 , 131 – 141 . support a branch containing the Guadalupe, South American, and Tedman , R. A. , and Bryden , M. M. ( 1981 ). The mammary-gland of the New Zealand fur seals and the Australian and New Zealand sea lion. Weddell seal, Leptonychotes weddelli (Pinnipedia) 1. Gross and Good support was also found for South African fur seals as sister microscopic anatomy . Anat. Rec. 199 , 519 – 529 . to this clade. The relationship between Northern ϩ California sea P Worthy , G. A. J. ( 1991 ). Insulation and thermal balance of fasting harp lions, Southern sea lion and remaining sea lions and Arctocephalus and gray seal pups . Comp. Biochem. Physiol. A 100 , 845 – 851 . species is not conclusively resolved. In support of fur seal and sea lion paraphyly is evidence for hybridization of various sympatric spe- cies (e.g., Arctocephalus gazella / A. tropicalis / A. forsteri , Zalophus californianus / Otaria byronia). Additionally, the violent sexual behav- ior by male sea lions toward females of different species my have resulted in more hybridization and introgression than has been typi- Pinnipedia, Overview cally recognized for the evolutionary history of otariids. ANNALISA BERTA B. Odobenidae: Walruses innipeds have always been understood to represent a group Although tusks are arguably the most characteristic feature of the distinct from other aquatic mammals. They are recog- modern walrus a rapidly improving fossil record indicates that these Pnized as members of the mammalian order Carnivora and unique structures evolved in a single lineage of walruses and “ tusks do include three monophyletic lineages, Otariidae (fur seals and sea not a walrus make ” ( Repenning and Tedford, 1977). The living walrus lions), Odobenidae (walruses), and Phocidae (true or earless seals). is the sole survivor of what was once a diverse radiation of at least 11 (A) (B) Figure 1 Representative otariids (A) southern sea lion, Otaria byronia and (B) South African fur seal, Arctocephalus pusillus, illustrating pinna. Note also the thick, dense fur characteristic of fur seals. Illustrations by P. Folkens. (From Berta et al., 2006 .) Figure 2 Distribution of sea lions. Based on Riedman (1990) . (From Berta et al., 2006 .) P Figure 3 Distribution of fur seals. Based on Riedman (1990) . (From Berta et al., 2006 .) 880 Pinnipedia, Overview genera and 14 species of walruses that lived from the early Miocene to molecular studies provide consistent, robust support for an alternative the end of the Pliocene (see also pinniped evolution ). Two subspe- alliance between otariids and odobenids. A survey of genetic variation cies of the modern walrus Odobenus rosmarus are usually recognized: among Atlantic and Pacifi c populations of the walrus suggests separa- O. r. rosmarus (Atlantic walrus) and O. rosmarus divergens (Pacifi c tion of the subspecies about 500,000– 750,000 years ago, supporting walrus) although a population from the Laptev Sea has been described the suggestion that Odobenus evolved in the Pacifi c and reached the as a third subspecies O. rosmarus. lapteevi but this has not been sub- North Atlantic early in Pleistocene time (Cronin et al ., 1994 ). stantiated. Pacifi c walruses are more abundant, are larger, and have longer tusks than Atlantic walruses ( Fay, 1981 ). Walruses inhabit the C. Phocidae: Seals northern hemisphere in areas with pack ice over shallow water of the The second major grouping of seals, the phocids, often are referred continental shelf ( Fig. 5 ). Like phocids, walruses lack external ear to as the “ true ” or “ earless ” seals for their lack of visible ear pinnae, fl aps. A unique feature of members of the modern walrus lineage are a characteristic which readily distinguishes them from otariids as enlarged upper canine tusks that function primarily in breeding and well as the walrus ( Fig. 6 ). Among the most distinguishing phocid social contexts. Walrus locomotion combines elements of phocid and otariid locomotion (discussed further under Adaptations). Monophyly
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