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Eared seal
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| Eared seals | |
|---|---|
| An Australian sea lion (Neophoca cinerea) | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Carnivora |
| Clade: | Panotariidae |
| Family: | Otariidae Gray, 1825 |
| Type genus | |
| Otaria Péron, 1816
| |
| Genera | |
An eared seal, otariid, or otary, is any member of the marine mammal family Otariidae, one of three groupings of pinnipeds. They comprise 15 extant species in seven genera (another species became extinct in the 1950s) and are commonly known either as sea lions or fur seals, distinct from true seals (phocids) and the walrus (odobenids). Otariids are adapted to a semiaquatic lifestyle, feeding and migrating in the water, but breeding and resting on land or ice. They reside in subpolar, temperate, and equatorial waters throughout the Pacific and Southern Oceans, the southern Indian, and Atlantic Oceans. They are conspicuously absent in the north Atlantic.
The words "otariid" and "otary" come from the Ancient Greek ὠτάριον (ōtárion), meaning "little ear",[2] referring to the small but visible external ear flaps (pinnae), which distinguishes them from the phocids.
Evolution and taxonomy
[edit]Morphological and molecular evidence supports a monophyletic origin of pinnipeds, sharing a common ancestor with Musteloidea,[3][4] though an earlier hypothesis suggested that Otаriidae are descended from a common ancestor most closely related to modern bears.[5] Debate remains as to whether the phocids diverged from the otariids before or after the walrus.
Otariids arose in the Miocene (15–17 million years ago) in the North Pacific, diversifying rapidly into the Southern Hemisphere, where most species now live. The earliest known fossil otariid is Eotaria crypta from southern California,[1] while the genus Callorhinus (northern fur seal) has the oldest fossil record of any living otariid, extending to the middle Pliocene. It probably arose from the extinct fur seal genus Thalassoleon.
Traditionally, otariids had been subdivided into the fur seal (Arctocephalinae) and sea lion (Otariinae) subfamilies, with the major distinction between them being the presence of a thick underfur layer in the former. Under this categorization, the fur seals comprised two genera: Callorhinus in the North Pacific with a single representative, the northern fur seal (C. ursinus), and eight species in the Southern Hemisphere under the genus Arctocephalus; while the sea lions comprise five species under five genera.[6] Recent analyses of the genetic evidence suggests that Callorhinus ursinus is in fact more closely related to several sea lion species.[7] Furthermore, many of the Otariinae appear to be more phylogenetically distinct than previously assumed; for example, the Japanese sea lion (Zalophus japonicus) is now considered a separate species, rather than a subspecies of the California sea lion (Zalophus californius).
In light of this evidence, the subfamily separation has been removed entirely and the family Otariidae has been organized into seven genera with 16 species and two subspecies.[8][9] Nonetheless, because of morphological and behavioral similarities among the "fur seals" and "sea lions", these remain useful categories when discussing differences between groups of species. Compared to sea lions, fur seals are generally smaller, exhibit greater sexual dimorphism, eat smaller prey and go on longer foraging trips; and, of course, there is the contrast between the coarse short sea lion hair and the fur seal's fur.
Cladogram showing relationships among the otarids, combining several phylogenetic analyses. The fur seal branch has not been resolved.[10][clarification needed]
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Anatomy
[edit]Otariids have proportionately much larger fore-flippers and pectoral muscles than phocids, and have the ability to turn their hind limbs forward and walk on all fours, making them far more maneuverable on land.[citation needed] They are generally considered to be less adapted to an aquatic lifestyle, since they breed primarily on land and haul out more frequently than true seals. However, they can attain higher bursts of speed and have greater maneuverability in the water. Their swimming power derives from the use of flippers more so than the sinuous whole-body movements typical of phocids and walruses.[citation needed]
Otariids are further distinguished by a more dog-like head, sharp, well-developed canines, and the aforementioned visible external pinnae. Their postcanine teeth are generally simple and conical in shape. The dental formula for eared seals is: 3.1.4.1-32.1.4.1. Sea lions are covered with coarse guard hairs, while fur seals have a thick underfur, which has historically made them the objects of commercial exploitation.[citation needed]
Male otariids range in size from the 70 kg (150 lb) Galápagos fur seal, smallest of all otariids, to the over 1,000-kg (2,200-lb) Steller sea lion. Mature male otariids weigh two to six times as much as females, with proportionately larger heads, necks, and chests, making them the most sexually dimorphic of all mammals.[11]
Behavior
[edit]All otariids breed on land during well-defined breeding seasons. Except for the Australian sea lion, which has an atypical 17.5 month breeding cycle, they form strictly annual aggregations on beaches or rocky substrates, often on islands. All species are polygynous; i.e. successful males breed with several females. In most species, males arrive at breeding sites first and establish and maintain territories through vocal and visual displays and occasional fighting. Females typically arrive on shore a day or so before giving birth. While considered social animals, no permanent hierarchies or statuses are established on the colonies. The extent to which males control females or territories varies between species. Thus, the northern fur seal and the South American sea lion tend to herd specific harem-associated females, occasionally injuring them, while the Steller sea lion and the New Zealand sea lion control spatial territories, but do not generally interfere with the movement of the females. Female New Zealand sea lions are the only otariids that move up to 2 km (1.2 mi) into forests to protect their pups during the breeding season.[12][13][14][15]
Otariids are carnivorous, feeding on fish, squid and krill. Sea lions tend to feed closer to shore in upwelling zones, feeding on larger fish, while the smaller fur seals tend to take longer, offshore foraging trips and can subsist on large numbers of smaller prey items. They are visual feeders. Some females are capable of dives of up to 400 m (1,300 ft).
Species
[edit]Family Otariidae
- Subfamily Arctocephalinae (fur seals)
- Genus Arctocephalus
- Brown fur seal, A. pusillus
- South African fur seal, A. pusillus pusillus
- Australian fur seal, A. pusillus doriferus
- Antarctic fur seal, A. gazella
- Guadalupe fur seal, A. townsendi
- Juan Fernández fur seal, A. philippii
- Galápagos fur seal, A. galapagoensis
- New Zealand fur seal (or southern fur seal), A. forsteri
- Subantarctic fur seal, A. tropicalis
- South American fur seal, A. australis
- Brown fur seal, A. pusillus
- Genus Callorhinus
- Northern fur seal, C. ursinus
- Genus Arctocephalus
- Subfamily Otariinae (sea lions)
- Genus Eumetopias
- Steller sea lion, E. jubatus
- Genus Neophoca
- Australian sea lion, N. cinerea
- Genus Otaria
- South American sea lion, O. flavescens
- Genus Phocarctos
- New Zealand sea lion (or Hooker's sea lion), P. hookeri
- Genus Zalophus
- California sea lion, Z. californianus
- †Japanese sea lion, Z. japonicus – extinct (1970s)[16][17]
- Galápagos sea lion, Z. wollebaeki
- Genus Eumetopias
Although the two subfamilies of otariids, the Otariinae (sea lions) and Arctocephalinae (fur seals), are still widely used, recent molecular studies have demonstrated that they may be invalid as traditionally defined.[18][19] Instead, they suggest three clades within the family; one consisting of the northern sea lions (Eumetopias and Zalophus), one of the northern fur seal (Callorhinus) and its extinct relatives, and the third of all the remaining Southern Hemisphere species.[20]
References
[edit]- ^ a b Boessenecker, R. W.; Churchill, M. (2015). "The oldest known fur seal". Biology Letters. 11 (2) 20140835. doi:10.1098/rsbl.2014.0835. PMC 4360102. PMID 25672999.
- ^ "Otary, n., etymology of" The Oxford English Dictionary. 2nd ed. 1989. OED Online. Oxford University Press. http://dictionary.oed.com/ Archived 2006-06-25 at the Wayback Machine Accessed November 2007
- ^ Hassanin, Alexandre; Veron, Géraldine; Ropiquet, Anne; Vuuren, Bettine Jansen van; Lécu, Alexis; Goodman, Steven M.; Haider, Jibran; Nguyen, Trung Thanh (2021-02-16). "Evolutionary history of Carnivora (Mammalia, Laurasiatheria) inferred from mitochondrial genomes". PLOS ONE. 16 (2) e0240770. Bibcode:2021PLoSO..1640770H. doi:10.1371/journal.pone.0240770. ISSN 1932-6203. PMC 7886153. PMID 33591975.
- ^ Berta, Annalisa (2009). "Pinniped Evolution". Encyclopedia of Marine Mammals. Elsevier. pp. 861–868. doi:10.1016/b978-0-12-373553-9.00199-1. ISBN 978-0-12-373553-9.
- ^ Lento, G.M.; Hickson, R.E.; Chambers, G.K.; Penny, D. (1 January 1995). "Use of spectral analysis to test hypotheses on the origin of pinnipeds". Molecular Biology and Evolution. 12 (1): 28–52. doi:10.1093/oxfordjournals.molbev.a040189. PMID 7877495.
- ^ J.E. King (1983). Seals of the World (2nd ed.). New York: Cornell University Press. ISBN 978-0-7022-1694-7.
- ^ Wynen, L; Goldsworthy, SD; Insley, SJ; Adams, M; Bickham, JW; Francis, J; Gallo, JP; Hoelzel, AR; et al. (2001). "Phylogenetic relationships within the eared seals (Otariidae: Carnivora): implications for the historical biogeography of the family". Mol. Phylogenet. Evol. 21 (2): 270–284. Bibcode:2001MolPE..21..270W. doi:10.1006/mpev.2001.1012. PMID 11697921.
- ^ Brunner, S. (2003). "Fur seals and sea lions (Otariidae): identification of species and taxonomic review". Systematics and Biodiversity. 1 (3): 339–439. doi:10.1017/S147720000300121X. S2CID 86375627.
- ^ "Otariidae". Integrated Taxonomic Information System.
- ^ Berta, Annalisa; Churchill, Morgan; Boessenecker, Robert W. (2018-05-30). "The Origin and Evolutionary Biology of Pinnipeds: Seals, Sea Lions, and Walruses". Annual Review of Earth and Planetary Sciences. 46 (1). Annual Reviews: 203–228. Bibcode:2018AREPS..46..203B. doi:10.1146/annurev-earth-082517-010009. ISSN 0084-6597. S2CID 135439365.
- ^ Weckerly, FW (1998). "Sexual-size dimorphism: influence of mass and mating systems in the most dimorphic mammals". Journal of Mammalogy. 79 (1): 33–42. doi:10.2307/1382840. JSTOR 1382840.
- ^ Augé, Amélie; Chilvers; Moore; Mathieu; Robertson (2009-01-01). "Aggregation and dispersion of female New Zealand sea lions at the Sandy Bay breeding colony, Auckland Islands: How unusual is their spatial behaviour?". Behaviour. 146 (9): 1287–1311. doi:10.1163/15683909X427687. ISSN 0005-7959.
- ^ Augé, Amélie A.; Chilvers, B. Louise; Mathieu, Renaud; Moore, Antoni B. (2012). "On-land habitat preferences of female New Zealand sea lions at Sandy Bay, Auckland Islands". Marine Mammal Science. 28 (3): 620–637. Bibcode:2012MMamS..28..620A. doi:10.1111/j.1748-7692.2011.00515.x. ISSN 1748-7692.
- ^ Frans, Veronica F.; Augé, Amélie A.; Edelhoff, Hendrik; Erasmi, Stefan; Balkenhol, Niko; Engler, Jan O. (2018). "Quantifying apart what belongs together: A multi-state species distribution modelling framework for species using distinct habitats". Methods in Ecology and Evolution. 9 (1): 98–108. Bibcode:2018MEcEv...9...98F. doi:10.1111/2041-210X.12847. ISSN 2041-210X. S2CID 91050320.
- ^ Frans, Veronica F.; Augé, Amélie A.; Fyfe, Jim; Zhang, Yuqian; McNally, Nathan; Edelhoff, Hendrik; Balkenhol, Niko; Engler, Jan O. (2022). "Integrated SDM database: Enhancing the relevance and utility of species distribution models in conservation management". Methods in Ecology and Evolution. 13 (1): 243–261. Bibcode:2022MEcEv..13..243F. doi:10.1111/2041-210X.13736. ISSN 2041-210X. S2CID 243893898.
- ^ Lowry, L. (2017). "Zalophus japonicus". IUCN Red List of Threatened Species. 2017 e.T41667A113089431. doi:10.2305/IUCN.UK.2017-1.RLTS.T41667A113089431.en. Retrieved 13 November 2021.
- ^ (in Japanese) Zalophus californianus japonicus (CR), Red Data Book, Japan Integrated Biodiversity Information System, Ministry of the Environment (Japan). "The Japanese sea lion (Zalophus californianus japonicus) was common in the past around the coast of the Japanese Archipelago, but declined rapidly after the 1930s from overhunting and increased competition with commercial fisheries. The last record in Japan was a juvenile, captured in 1974 off the coast of Rebun Island, northern Hokkaido."
- ^ Yonezawa, T.; et al. (2009). "The monophyletic origin of sea lions and fur seals (Carnivora; Otariidae) in the Southern Hemisphere". Gene. 441 (1–2): 89–99. doi:10.1016/j.gene.2009.01.022. PMID 19254754.
- ^ Arnason, U.; et al. (2006). "Pinniped phylogeny and a new hypothesis for their origin and dispersal". Molecular Phylogenetics and Evolution. 41 (2): 345–354. Bibcode:2006MolPE..41..345A. doi:10.1016/j.ympev.2006.05.022. PMID 16815048.
- ^ Berta, A.; Churchill, M. (2012). "Pinniped Taxonomy: evidence for species and subspecies". Mammal Review. 42 (3): 207–234. doi:10.1111/j.1365-2907.2011.00193.x.
Further reading
[edit]- Berta, A., and L. Sumich (1999) Marine Mammals: Evolutionary Biology. San Diego: Academic Press.
- Gentry, R. L (1998) Behavior and Ecology of the Northern Fur Seal. Princeton: Princeton University Press.
- Perrin, W. F., B. Würsig, and J. G. M. Thewissen (2002) Encyclopedia of Marine Mammals. San Diego: Academic Press.
- Riedman, M. (1990) The Pinnipeds: Seals, Sea Lions and Walruses. Berkeley: University of California Press.
External links
[edit]Eared seal
View on GrokipediaTaxonomy and evolution
Taxonomy
Eared seals constitute the family Otariidae, the only family in the superfamily Otarioidea, which is part of the suborder Pinnipedia within the order Carnivora.[6] This classification places Otariidae alongside the families Phocidae (true seals) and Odobenidae (walruses) as one of the three extant pinniped families.[7] The current taxonomy recognizes 7 genera and 14 extant species in Otariidae.[8][1][9] The genera include Arctocephalus (southern fur seals), Callorhinus (northern fur seal), Eumetopias (Steller sea lion), Neophoca (Australian sea lion), Otaria (South American sea lion), Phocarctos (New Zealand sea lion), and Zalophus (California and Galápagos sea lions).[1] Historically, Otariidae was divided into two subfamilies: Arctocephalinae for fur seals (genera Arctocephalus and Callorhinus), distinguished by their dense underfur, and Otariinae for sea lions (the remaining genera), which lack such underfur. However, molecular phylogenetic analyses have challenged this bipartition, demonstrating that fur seals form a paraphyletic group and that the family comprises three distinct clades: one uniting the northern fur seal (Callorhinus) with California sea lions (Zalophus), another including the Steller sea lion (Eumetopias), and a third encompassing the southern fur seals (Arctocephalus) and other southern sea lions.[7] Otariidae represents a monophyletic group that diverged from the common ancestor shared with Phocidae and Odobenidae approximately 20-25 million years ago.[7]Evolutionary history
The eared seals (family Otariidae) originated in the North Pacific Ocean during the middle Miocene epoch, approximately 15–17 million years ago (Ma), evolving from an 'enaliarctine'-like ancestor that represented early pinnipedimorphs transitioning from terrestrial carnivorans to marine lifestyles.[10] The earliest known otariid fossil, Eotaria crypta, dated to 14.9–17.1 Ma from the Topanga Formation in southern California, exhibits morphological intermediates such as a shortened rostrum and specialized dentition adapted for pelagic feeding, filling a previous geochronological gap and eliminating a ghost lineage of about 5–6 Ma between stem pinnipedimorphs and crown otariids.[11] Earlier transitional forms, including Pteronarctos from the late early Miocene (approximately 20–15 Ma) in Oregon, show key adaptations like enlarged orbits and reduced limbs, bridging arctoid mustelids to fully aquatic pinnipeds, though Pteronarctos itself lies outside the crown Pinnipedia clade.[12] Phylogenetically, Otariidae forms part of the monophyletic order Pinnipedia, with molecular and morphological evidence resolving earlier debates over diphyly (separate origins for otariids/odobenids from phocids) in favor of a single origin from a common arctoid ancestor closely related to ursids.[12] Within Pinnipedia, Otariidae is the sister group to Odobenidae (walruses), with this Otarioidea clade sister to Phocidae (true seals), as supported by mitochondrial and nuclear DNA analyses that place the basal divergence of otariids around 16 Ma.[13] This monophyly is reinforced by shared morphological and molecular synapomorphies, contrasting with the earlier paraphyletic hypotheses based solely on morphology.[14] Diversification of Otariidae accelerated in the late Miocene to early Pliocene (approximately 10–5 Ma), with radiation into subfamilies Arctocephalinae (fur seals) and Otariinae (sea lions) driven by cooling climates, changing ocean currents, and habitat expansion.[15] Early otariids like Pithanotaria (late Miocene, ~11 Ma, California) represent basal fur seal-like forms, while later fossils such as Thalassoleon (late Miocene to early Pliocene) indicate sea lion divergence.[12] A key event was the dispersal to the Southern Hemisphere around 6 Ma, facilitated by the Antarctic Circumpolar Current, which enabled southward migration along western South America and subsequent isolation leading to endemic radiations in subantarctic waters.[15] This vicariance, combined with Northern Hemisphere retention of lineages like northern fur seals, shaped the family's current bipolar distribution.[16]Physical characteristics
Anatomy
Eared seals, members of the family Otariidae, exhibit a suite of anatomical adaptations that facilitate their semiaquatic lifestyle, balancing efficient propulsion in water with mobility on land. Their skeletal framework supports powerful swimming via modified limbs, while sensory organs are tuned for both aerial and aquatic environments. Physiological modifications in respiration and circulation enable extended submersion, and specialized cranial features aid in prey processing. Unlike true seals (Phocidae), which rely heavily on blubber, fur seals emphasize dense fur for insulation while sea lions depend more on blubber layers.[17] The skeletal structure of eared seals is characterized by elongated forelimbs that form broad, paddle-like flippers essential for aquatic propulsion. These foreflippers display hyperphalangy, an increase in phalangeal elements beyond the typical mammalian count, which enhances flexibility and surface area for maneuvering in water.[18] In contrast, the hind flippers are uniquely rotatable, allowing them to pivot forward under the body for a walking gait on land, a capability absent in true seals that instead undulate to move.[19] This rotation is facilitated by a flexible pelvic girdle, enabling more agile terrestrial locomotion compared to the belly-crawling of earless seals.[20] Sensory systems in eared seals are highly specialized for detecting stimuli in varied media. Prominent external ear pinnae, or auricles, collect sound waves effectively in air, contributing to acute hearing, with underwater sensitivity peaking in the 15–30 kHz range for species like the California sea lion.[21] Vibrissae, or whiskers, are richly innervated with up to 1,500 nerve endings per follicle, serving as mechanoreceptors to detect hydrodynamic trails from prey during underwater navigation.[22] Their eyes feature a tapetum lucidum, a reflective layer behind the retina that amplifies low-light vision by redirecting photons through rod cells, aiding detection in dimly lit aquatic environments.[21] Respiratory and circulatory adaptations support diving capabilities typical of eared seals, with typical dives lasting 1–5 minutes and maximum durations up to 10–15 minutes. High concentrations of myoglobin in skeletal muscles store oxygen efficiently, allowing reliance on anaerobic metabolism during submersion and reducing the need for frequent surfacing.[23] Circulatory systems incorporate countercurrent heat exchangers in the flippers, where arterial blood warms venous return to conserve core body heat in cold waters, preventing hypothermia without excessive energy expenditure.[24] These vascular networks minimize heat loss to the periphery, particularly important given the thinner blubber layer compared to true seals.[3] Cranial anatomy includes a dog-like skull with pronounced carnassial teeth—specialized upper and lower molars that shear against each other for efficient prey dismemberment.[25] This dentition reflects their carnivorous diet, with sharp canines for grasping and postcanine teeth adapted for tearing fish and squid. For insulation, eared seals possess dense, multilayered fur that traps air for thermal protection, supplemented by a modest blubber layer, differing from the thick blubber-dominated insulation of true seals.[26] This fur-based system supports their more active terrestrial behaviors while maintaining buoyancy and streamlining in water.[17]Size, appearance, and dimorphism
Eared seals exhibit a wide range of body sizes across the Otariidae family, with the smallest species being the Galápagos fur seal (Arctocephalus galapagoensis), where adult males reach lengths of about 1.5 m and weights of up to 64 kg, while females are smaller at around 1.2 m and 27 kg on average.[27] In contrast, the largest species, the Steller sea lion (Eumetopias jubatus), features adult males that can grow to over 3 m in length and exceed 1,000 kg in weight, whereas females typically measure 2.3–2.9 m and weigh up to 363 kg.[28] These size variations reflect adaptations to diverse marine environments, but all eared seals share a streamlined, torpedo-shaped body that facilitates efficient swimming.[2] The external appearance of eared seals is characterized by a sleek, elongate form covered in coarse guard hairs overlying a dense underfur, which provides insulation and waterproofing, particularly in fur seal species. Coloration differs notably between fur seals and sea lions: fur seals often display a silvery-gray sheen when dry due to their thick, fine underfur, while sea lions tend toward tawny brown or dark tones, with males darkening to nearly black in some cases like the California sea lion (Zalophus californianus).[29] External ear flaps, a defining trait of the family, are visible and aid in thermoregulation on land.[17] Sexual dimorphism in eared seals is pronounced, with adult males generally 2–5 times heavier than females, enabling males to dominate breeding territories but requiring females to be more agile for foraging and pup-rearing. Males often develop broader heads, thicker necks, and in species like the California sea lion, a prominent sagittal crest that gives a maned appearance, enhancing their competitive displays during the breeding season. Females, by comparison, maintain a slimmer, more hydrodynamic build suited to extended dives.[30][31] Fur seals undergo an annual molting cycle, shedding their dense underfur in a rapid process that lasts several weeks, typically in late summer or fall, which temporarily reduces insulation and increases vulnerability to heat loss and predation as they haul out onshore without feeding. This molt replaces the underfur essential for buoyancy and thermal protection, distinguishing fur seals from sea lions, which undergo a slower, less disruptive hair replacement.[32]Distribution and habitat
Geographic range
Eared seals, members of the family Otariidae, exhibit a predominantly Northern Hemisphere distribution centered in the North Pacific Ocean, with significant extensions into the Southern Hemisphere along temperate and subpolar coasts. Most species inhabit coastal and pelagic waters of the Pacific, including subarctic regions, but they are absent from true Arctic environments where breeding populations of true seals (Phocidae) dominate. Their range spans from equatorial waters near the Galápagos Islands to sub-Antarctic islands, reflecting adaptations to a variety of marine ecosystems without crossing into polar ice-dominated Arctic habitats.[17][33] In the Northern Hemisphere, key species like the northern fur seal (Callorhinus ursinus) are distributed across the Bering Sea, Sea of Okhotsk, Sea of Japan, and southward to the California coast, with primary breeding colonies on islands such as the Pribilofs and Commander Islands. The Steller sea lion (Eumetopias jubatus) occupies a broad North Pacific range from central California northward through Alaska and the Aleutian Islands to the Kuril Islands and Japan, favoring rocky shores and coastal waters. These distributions highlight the family's concentration in productive North Pacific upwelling zones.[4][34][35][36] Southern Hemisphere extensions include the South American sea lion (Otaria flavescens), which ranges along the Pacific and Atlantic coasts from northern Peru southward to Cape Horn and eastward to Uruguay and southern Brazil. The Antarctic fur seal (Arctocephalus gazella) is centered on sub-Antarctic islands south of the Antarctic Convergence, with over 95% of its breeding population at South Georgia and nearby sites like the South Shetlands. The Galápagos fur seal (Arctocephalus galapagoensis) is endemic to the Galápagos Archipelago off Ecuador, occurring primarily on western islands in equatorial Pacific waters. In the Indian Ocean, the South African fur seal (Arctocephalus pusillus) inhabits coasts from Angola to Mozambique, including islands like the Robben Islands, while the Australian sea lion (Neophoca cinerea) is restricted to waters off southern and western Australia.[37][38][39][40][27][2] While eared seals maintain stable core ranges, vagrant individuals occasionally appear in unexpected areas, such as Antarctic fur seals recorded off the eastern Atlantic islands like Tristan da Cunha. No established populations exist in the Arctic, though rare sightings of northern species have been noted near Arctic fringes without breeding. These vagrancies underscore the species' dispersive nature but do not alter their primary temperate and subpolar distributions.[41]Habitat preferences
Eared seals primarily utilize terrestrial habitats consisting of rocky shores, sandy beaches, and isolated islands for haul-outs, breeding rookeries, and molting sites, favoring remote locations that minimize predation risks.[4] These sites provide essential resting and thermoregulatory opportunities, with individuals often aggregating in large colonies during breeding seasons to facilitate social interactions and pup rearing.[20] Preference for such predator-free environments enhances survival rates for vulnerable pups and nursing females.[42] In aquatic environments, eared seals occupy coastal and pelagic waters across temperate, subpolar, and equatorial regions with a wide range of sea surface temperatures, from about 5°C in subpolar waters to 25°C or higher in equatorial regions, which support their metabolic and foraging needs.[24][27] They are adapted to dive to depths of 200–500 meters during foraging excursions, though most activity occurs in shallower coastal zones up to 100–300 meters.[43] These habitats allow for efficient navigation between land-based resting areas and offshore prey resources.[44] Fur seals exhibit a stronger affinity for offshore upwelling zones, where nutrient-rich waters promote high prey densities such as fish and squid, enabling pelagic foraging strategies.[25] In contrast, sea lions tend to remain nearer to shorelines, exploiting nearshore currents and benthic habitats for more localized hunting.[45] This divergence in habitat use reflects differences in body insulation and locomotion, with fur seals relying on dense pelage for extended open-water travel.[46] Breeding colonies are typically established on steep cliffs or expansive sandy beaches that offer protection from waves and terrestrial predators, while sensitivity to oceanographic features like upwelling currents influences site selection for prey proximity.[47] These microhabitats ensure accessibility for haul-outs and support the energetic demands of reproduction in dynamic marine ecosystems.[48]Behavior and ecology
Social structure and reproduction
Eared seals, or otariids, typically form large colonies known as rookeries on beaches or rocky shores during the breeding season, where social interactions are structured around dominance hierarchies among males.[49] Dominant adult males establish and defend territories through aggressive displays, vocalizations, and physical confrontations, often lasting several weeks to secure access to females.[50] These hierarchies reduce the need for constant fighting by signaling status, with subordinate males relegated to peripheral or bachelor groups.[51] The mating system of eared seals is predominantly polygynous, characterized by resource-defense where dominant males, termed beachmasters, control territories and herd harems of 10 to 40 females to monopolize mating opportunities.[50] However, genetic studies reveal variability in reproductive skew; for instance, in California sea lions, only a subset of territorial males sire offspring, with most fathering one or two pups rather than large numbers, indicating weaker polygyny than behavioral observations suggest.[52] Extra-territorial copulations are common, particularly during female foraging trips at sea, allowing subordinate or non-territorial males to achieve paternity outside defended areas.[52] Reproduction in eared seals is seasonal, with northern hemisphere species breeding from May to August and southern hemisphere species from November to February, aligning with optimal environmental conditions for pup rearing.[50] Gestation lasts 8 to 12 months, incorporating delayed implantation where the fertilized embryo remains unattached in the uterus for 2 to 4 months before development resumes, ensuring births coincide with abundant food resources.[53] Pups are born precocial, able to move and swim shortly after birth, and are nursed for 4 to 12 months, during which females alternate between foraging at sea and returning to feed their young.[50] Notably, the Australian sea lion exhibits an asynchronous breeding cycle of 17.5 months across its range, differing from the annual patterns in most otariids and resulting in less frequent pup production.[54] Infanticide rates are elevated in some eared seal species following male territorial takeovers, as incoming dominant males kill unrelated pups to hasten female estrus and redirect reproductive effort toward their own offspring.[55] In the New Zealand sea lion, for example, such behavior has been observed among non-territorial males, often linked to energy acquisition through subsequent cannibalism.[55]Foraging and diet
Eared seals, members of the family Otariidae, exhibit primarily piscivorous and cephalopod-based diets, with prey selection varying between fur seals and sea lions based on habitat and foraging mode. Fur seals, such as the northern fur seal (Callorhinus ursinus) and Antarctic fur seal (Arctocephalus gazella), predominantly consume pelagic squid and small schooling fish like lanternfish and sardines, often targeting these resources nocturnally in open ocean waters.[4][44] In contrast, sea lions, including the California sea lion (Zalophus californianus) and South American sea lion (Otaria flavescens), focus on nearshore fish such as herring, sardines, rockfish, and hake, supplemented by cephalopods like market squid and occasional crustaceans or octopus.[29] Some species incorporate krill during periods of high availability, particularly in subantarctic regions.[56] Foraging occurs solitarily at sea, with individuals employing breath-hold dives lasting 1–10 minutes to depths of 50–700 meters, depending on species and prey distribution; fur seals typically engage in shallower, shorter epi- and mesopelagic dives (e.g., 10–60 m for 0.5–3 minutes in northern fur seals), while certain sea lions, like the New Zealand sea lion (Phocarctos hookeri), reach benthic depths up to 600 m for up to 20 minutes.[56][4] They detect prey using specialized vibrissae (whiskers) that sense hydrodynamic wakes from swimming fish or squid, enabling tracking in low-visibility conditions without echolocation.[57] Foraging strategies shift seasonally with prey availability; for instance, California sea lions consume more sardines and anchovies in summer upwelling periods but diversify to squid and rockfish in winter. Lactating northern fur seals ingest approximately 9.5–10 kg of prey daily during foraging trips to support energy demands. As apex predators, eared seals regulate fish and cephalopod populations, exerting top-down control on marine food webs and influencing community structure through selective predation on schooling species.[56] Their consumption overlaps with commercial fisheries, leading to resource competition; for example, northern fur seals target walleye pollock, a key fishery species in the Bering Sea, potentially amplifying pressures on shared stocks during periods of low prey abundance.[4] Juveniles acquire foraging skills through observation and accompaniment on maternal trips post-weaning, gradually developing independent diving and prey-capture abilities over the first year.[58]Conservation and interactions
Threats and status
Eared seals face a range of conservation challenges, with IUCN Red List assessments indicating that out of 15 extant species, three are classified as Endangered (the New Zealand sea lion Phocarctos hookeri, Galápagos sea lion Zalophus wollebaeki, and Australian sea lion Neophoca cinerea), the Galápagos fur seal Arctocephalus galapagoensis as Endangered, the Guadalupe fur seal Arctocephalus townsendi and Juan Fernández fur seal Arctocephalus philippii as Least Concern (as of 2023), the Japanese sea lion Zalophus japonicus as Extinct (last confirmed in the 1970s), and the remainder as Least Concern or Near Threatened (e.g., Steller sea lion Eumetopias jubatus as Near Threatened globally, though its western population is Endangered under U.S. law).[59][60][61] Primary threats include incidental capture in commercial fishing gear, climate-induced shifts in prey availability, and persistent pollutants. Bycatch remains a significant mortality factor, particularly for larger species; for instance, estimates from the 1980s indicated over 1,000 Steller sea lions caught annually in the western Bering Sea trawl fisheries. Climate change exacerbates vulnerability by altering ocean temperatures and prey distributions, leading to nutritional stress; historical overhunting in the 19th century reduced northern fur seal (Callorhinus ursinus) populations by up to 90% on the Pribilof Islands and southern fur seal (Arctocephalus spp.) populations, such as the Antarctic fur seal, by nearly 100% in subantarctic regions, contributing to long-term genetic bottlenecks that hinder resilience.[62][63][64] Population trends vary across species, with declines observed in at least five, including a greater than 70% reduction in Steller sea lion numbers since 1980, primarily in the Gulf of Alaska and Aleutian Islands due to combined fishery and environmental pressures; however, western populations have stabilized since the early 2000s. In contrast, some populations have shown recovery, such as the California sea lion Zalophus californianus, which has increased steadily following 20th-century protections, though unusual mortality events linked to domoic acid from harmful algal blooms affected pups in 2023-2024. Specific events like strong El Niño oscillations intensify starvation risks for equatorial species, causing pup mortality rates to spike as sardine and anchovy schools shift away from breeding grounds, as seen in the 2015–2016 event affecting California sea lions. Pollution from polychlorinated biphenyls (PCBs) has been linked to reproductive failures in otariids, including reduced fertility and increased miscarriage rates through endocrine disruption, with elevated levels documented in blubber samples from recovering populations.[65][66][29][67]Human relationships and conservation
Human interactions with eared seals have been marked by extensive exploitation, particularly through commercial fur hunting that began in the late 18th century and peaked during the 19th century, leading to severe population declines. On the Pribilof Islands in Alaska, northern fur seals (Callorhinus ursinus) were targeted intensively starting in 1786, with Russian and later American hunters harvesting hundreds of thousands of skins annually; by the mid-19th century, unregulated pelagic sealing had decimated breeding colonies, reducing populations from millions to critically low levels.[68][69] This overharvesting extended to other eared seal species, such as Antarctic fur seals (Arctocephalus gazella), where commercial operations in subantarctic regions slaughtered up to two million individuals between 1800 and 1820, nearly driving the species to extinction.[70] Indigenous communities have long incorporated eared seals into their cultural practices, utilizing them for sustenance and materials in sustainable ways predating European contact. In the North Pacific, Native American groups along the Northwest Coast, including tribes in present-day Washington and British Columbia, hunted northern fur seals for meat, oil, and hides, maintaining populations through selective harvesting of non-breeding males over millennia.[71] Similarly, Aleut peoples on the Pribilof Islands traditionally relied on fur seals for food, clothing from skins, and tools, a practice that continued under regulated subsistence quotas after commercial hunting ended.[72] In more recent times, ecotourism has highlighted eared seals' role in cultural and economic contexts, such as in the Galápagos Islands, where Galápagos sea lions (Zalophus wollebaeki) draw visitors through guided snorkeling and beach observations, supporting local economies while promoting awareness of marine biodiversity.[73] Conservation efforts for eared seals have evolved from international agreements to address historical overexploitation. The 1911 North Pacific Fur Seal Treaty, signed by the United States, Russia, Japan, and Great Britain (on behalf of Canada), prohibited open-sea hunting of northern fur seals and established shared management of rookeries, allowing populations to recover from near-extinction levels.[4] Protected areas have further bolstered recovery; for instance, Channel Islands National Park in California safeguards California sea lions (Zalophus californianus) through marine protected areas that restrict fishing and promote habitat integrity, contributing to stable or increasing colony sizes.[74][75] To mitigate ongoing fisheries interactions, bycatch reduction technologies such as top-opening excluder devices in trawl nets have been implemented, enabling pinnipeds like sea lions to escape gear and reducing incidental mortality in commercial operations.[76][77] Sustainable harvest management now focuses on subsistence needs with strict quotas to prevent overexploitation. For northern fur seals on the Pribilof Islands, annual subsistence harvests are limited to 1,645–2,000 juvenile males on St. Paul Island and up to 150 on St. George Island, monitored to ensure long-term viability without impacting breeding stocks.[78] Regarding extinct species, reintroduction of the Japanese sea lion (Zalophus japonicus), last sighted in the 1970s, has been proposed by entities including the South Korean Ministry of Environment, but efforts remain debated due to genetic and ecological uncertainties in sourcing surrogates from related taxa.[79]Species
Fur seals
Fur seals belong to two genera within the subfamily Arctocephalinae: Arctocephalus, which encompasses nine species of southern fur seals primarily distributed in the Southern Hemisphere, and Callorhinus, represented by a single species, the northern fur seal (C. ursinus), found in the North Pacific. A defining characteristic of fur seals is their dense underfur, consisting of up to 350,000 fine hairs per square inch overlaid by coarser guard hairs, which provides exceptional insulation in cold marine environments and historically drove intense commercial hunting for pelts beginning in the late 18th century.[33][4] This underfur, combined with their specialized foraging behavior, positions fur seals as offshore or pelagic hunters, often diving to depths exceeding 200 meters to pursue prey like squid, fish, and krill far from coastal zones, distinguishing them from more nearshore-oriented relatives.[80][81] The group exhibits remarkable species diversity, with Arctocephalus species ranging from temperate to subantarctic waters across oceans, including the South American fur seal (A. australis) along Pacific and Atlantic coasts, the New Zealand fur seal (A. forsteri) in Australasian waters, and the Guadalupe fur seal (A. townsendi) off western North America.[82] Among these, the Antarctic fur seal (A. gazella) stands out for its abundance, with a global population estimated between 1.5 and 4 million individuals concentrated around the Antarctic Peninsula and subantarctic islands, while the Galápagos fur seal (A. galapagoensis), the smallest species at about 1.2-1.5 meters in length and endemic to the Galápagos Islands, numbers around 10,000-15,000 and faces localized pressures from El Niño events.[83] All fur seal species display extreme sexual dimorphism, with adult males typically 2-5 times heavier and 20-40% longer than females, enabling males to defend breeding territories but limiting their agility during extended foraging trips.[38][84] Following near-extirpation from 19th-century overhunting, many populations have rebounded through international protections like the 1911 Fur Seal Treaty, yet they remain vulnerable to ocean warming, which disrupts prey availability—such as krill declines linked to rising sea temperatures—and has contributed to recent population stagnation or decreases in species like the Antarctic fur seal.[4][85] The following table summarizes key details for fur seal species:| Species | Scientific Name | Primary Range | IUCN Status | Population Estimate (approx., as of 2023) |
|---|---|---|---|---|
| Northern fur seal | Callorhinus ursinus | North Pacific (Alaska to California, Russia to Japan) | Vulnerable | 1,000,000–1,200,000 |
| Antarctic fur seal | Arctocephalus gazella | Antarctic and subantarctic islands | Least Concern | 2,000,000–4,000,000 |
| South American fur seal | Arctocephalus australis | South American coasts (Pacific and Atlantic) | Least Concern | 200,000–500,000 |
| New Zealand fur seal | Arctocephalus forsteri | Australia, New Zealand, subantarctic islands | Least Concern | 100,000–200,000 |
| Galápagos fur seal | Arctocephalus galapagoensis | Galápagos Islands | Vulnerable | 10,000–15,000 |
| Guadalupe fur seal | Arctocephalus townsendi | Baja California, Mexico | Least Concern | 20,000–30,000 |
| Juan Fernández fur seal | Arctocephalus philippii | Juan Fernández Islands, Chile | Vulnerable | 5,000–10,000 |
| Subantarctic fur seal | Arctocephalus tropicalis | Subtropical islands (e.g., Gough, Tristan da Cunha) | Least Concern | 200,000–300,000 |
| Cape fur seal | Arctocephalus pusillus | Southern Africa coasts and islands | Least Concern | 1,500,000–2,000,000 |
| Australian fur seal | Arctocephalus mackenzii | Southern Australia | Vulnerable | 20,000–30,000 |
Sea lions
Sea lions belong to the subfamily Otariinae of the family Otariidae, distinguishing them from the fur seals of the subfamily Arctocephalinae.[30] Unlike fur seals, sea lions lack dense underfur and instead rely on a thick layer of blubber for thermal insulation, while also exhibiting a more robust body build and a pronounced dog-like facial structure with a sloping forehead.[3] They share the family's defining traits, including visible external ear flaps (pinnae), elongated foreflippers for swimming propulsion, and rotatable hind flippers that enable terrestrial locomotion on all fours, setting them apart from true seals (Phocidae) which lack ear flaps and use hind flippers primarily for swimming.[2] Sexually dimorphic across all species, adult males are typically 1.5 to 2 times larger than females and often develop thick manes around the neck and shoulders during breeding season.[3] Sea lions are highly social marine mammals, often forming large colonies on rocky shores or beaches for breeding and resting, with vocalizations including loud barks and roars that facilitate communication in noisy rookeries.[3] They are agile swimmers, using a powerful "clap" motion of the foreflippers to reach speeds up to 40 km/h (25 mph) and capable of diving to depths of 100-300 meters for foraging, though they generally prefer coastal waters over open ocean.[29][87] Their diet consists mainly of fish, squid, and occasionally seabirds or crustaceans, hunted in groups or individually depending on prey availability.[28] Reproductive cycles vary by species but typically involve polygynous mating systems where dominant males defend harems of females on rookeries, with gestation lasting 8-12 months and pups nursed for several months to a year.[2] Six extant species of sea lions are recognized, distributed across temperate and subpolar coastal regions of the Pacific, southern, and South Atlantic oceans.[88] These species exhibit varying population sizes and conservation statuses, influenced by historical exploitation, fisheries bycatch, and habitat pressures. The following table summarizes key details:| Species | Scientific Name | Primary Range | IUCN Status | Population Estimate (approx., as of 2023) |
|---|---|---|---|---|
| California sea lion | Zalophus californianus | Western coast of North America, from Canada to Mexico | Least Concern | 180,000–300,000 |
| Galápagos sea lion | Zalophus wollebaeki | Galápagos Islands, Ecuador | Endangered | 10,000–20,000 |
| Steller sea lion | Eumetopias jubatus | North Pacific, from California to Japan | Near Threatened | 180,000–250,000 |
| South American sea lion | Otaria flavescens | Coasts of Peru, Chile, Argentina, Uruguay | Least Concern | 400,000–450,000 |
| Australian sea lion | Neophoca cinerea | Southern and western Australia | Endangered | 10,000–12,000 |
| New Zealand sea lion | Phocarctos hookeri | Subantarctic islands and mainland New Zealand | Endangered | 10,000–13,000 |