Seabirds longlinersm.jpg

Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same environmental problems and feeding niches have resulted in similar adaptations. The first seabirds evolved in the Cretaceous period, and modern seabird families emerged in the Paleogene.

In general, seabirds live longer, breed later and have fewer young than other birds do, but they invest a great deal of time in their young. Most species nest in colonies, which can vary in size from a few dozen birds to millions. Many species are famous for undertaking long annual migrations, crossing the equator or circumnavigating the Earth in some cases. They feed both at the ocean's surface and below it, and even feed on each other. Seabirds can be highly pelagic, coastal, or in some cases spend a part of the year away from the sea entirely.

Seabirds and humans have a long history together: they have provided food to hunters, guided fishermen to fishing stocks and led sailors to land. Many species are currently threatened by human activities, such as from oil spills, getting trapped in nets, and by climate change and severe weather. Conservation efforts include the establishment of wildlife refugees and adjustments to fishing techniques.

Classification

There exists no single definition of which groups, families and species are seabirds, and most definitions are in some way arbitrary. Elizabeth Shreiber and Joanne Burger, two seabird scientists, said, "The one common characteristic that all seabirds share is that they feed in saltwater; but, as seems to be true with any statement in biology, some do not."Schreiber, Elizabeth A. and Burger, Joanne (2001) ''Biology of Marine Birds''. Boca Raton: CRC Press, However, by convention all of the Sphenisciformes (penguins) and Procellariiformes ( albatrosses and petrels), all of the Suliformes (gannets and cormorants) except the darters, and some of the Charadriiformes (the skuas, gulls, terns, auks and skimmers) are classified as seabirds. The phalaropes are usually included as well, since although they are waders ("shorebirds" in North America), two of the three species (Red and Red-necked) are oceanic for nine months of the year, crossing the equator to feed pelagically.

Loons and grebes, which nest on lakes but winter at sea, are usually categorized as water birds, not seabirds. Although there are a number of sea ducks in the family Anatidae that are truly marine in the winter, by convention they are usually excluded from the seabird grouping. Many waders (or shorebirds) and herons are also highly marine, living on the sea's edge (coast), but are also not treated as seabirds. Sea eagles and other fish-eating birds of prey are also typically excluded, however tied to marine environments they may be.

German paleontologist Gerald Mayr defined the "core waterbird" clade Aequornithes in 2010. This lineage gives rise to the Gaviiformes, Sphenisciformes, Procellariiformes, Ciconiiformes, Suliformes and Pelecaniformes. The tropicbirds are part of a lineage—Eurypygimorphae—that is a sister group to the Aequornithes.

Evolution and fossil record

Seabirds, by virtue of living in a geologically depositional environment (that is, in the sea where sediments are readily laid down), are well represented in the fossil record. They are first known to occur in the Cretaceous period, the earliest being the Hesperornithiformes, like ''Hesperornis regalis'', a flightless loon-like seabird that could dive in a fashion similar to grebes and loons (using its feet to move underwater) but had a beak filled with sharp teeth. Flying Cretaceous seabirds do not exceed wingspans of two meters; any sizes were taken by piscivorous pterosaurs.

While ''Hesperornis'' is not thought to have left descendants, the earliest modern seabirds also occurred in the Cretaceous, with a species called '' Tytthostonyx glauconiticus'', which has features suggestive of Procellariiformes and Fregatidae.Olson, S.; Parris, D.C. (1987).
The Cretaceous Birds of New Jersey.
''Smithsonian Contributions to Paleontology'', 63: 22pp. As a clade, the Aequornithes either became seabirds in a single transition in the Cretaceous or some lineages such as pelicans and frigatebirds adapted to sea living independently from freshwater-dwelling ancestors. In the Paleogene both pterosaurs and marine reptiles became extinct, allowing seabirds to expand ecologically. These post-extinction seas were dominated by early Procellariidae, giant penguins and two extinct families, the Pelagornithidae and the Plotopteridae (a group of large seabirds that looked like the penguins). Modern genera began their wide radiation in the Miocene, although the genus ''Puffinus'' (which includes today's Manx shearwater and sooty shearwater) might date back to the Oligocene. Within the Charadriiformes, the gulls and allies ( Lari) became seabirds in the late Eocene, and then waders in the middle Miocene (Langhian). The highest diversity of seabirds apparently existed during the Late Miocene and the Pliocene had undergone a period of upheaval due to extinction of considerable numbers of marine species; subsequently, the spread of marine mammals seems to have prevented seabirds from reaching their erstwhile diversity.

Characteristics

Adaptations to life at sea

Seabirds have made numerous adaptations to living on and feeding in the sea. Wing morphology has been shaped by the niche an individual species or family has evolved, so that looking at a wing's shape and loading can tell a scientist about its life feeding behaviour. Longer wings and low wing loading are typical of more pelagic species, while diving species have shorter wings.Gaston, Anthony J. (2004). ''Seabirds: A Natural History'' New Haven:Yale University Press, Species such as the wandering albatross, which forage over huge areas of sea, have a reduced capacity for powered flight and are dependent on a type of gliding called dynamic soaring (where the wind deflected by waves provides lift) as well as slope soaring. Seabirds also almost always have webbed feet, to aid movement on the surface as well as assisting diving in some species. The Procellariiformes are unusual among birds in having a strong sense of smell, which is used to find widely distributed food in a vast ocean, and help distinguish familiar nest odours from unfamiliar ones.

Salt glands are used by seabirds to deal with the salt they ingest by drinking and feeding (particularly on crustaceans), and to help them osmoregulate.Harrison, C. S. (1990) ''Seabirds of Hawaii, Natural History and Conservation'' Ithaca:Cornell University Press, The excretions from these glands (which are positioned in the head of the birds, emerging from the nasal cavity) are almost pure sodium chloride.

With the exception of the cormorants and some terns, and in common with most other birds, all seabirds have waterproof plumage. However, compared to land birds, they have far more feathers protecting their bodies. This dense plumage is better able to protect the bird from getting wet, and cold is kept out by a dense layer of down feathers. The cormorants possess a layer of unique feathers that retain a smaller layer of air (compared to other diving birds) but otherwise soak up water. This allows them to swim without fighting the buoyancy that retaining air in the feathers causes, yet retain enough air to prevent the bird losing excessive heat through contact with water.

The plumage of most seabirds is less colourful than that of land birds, restricted in the main to variations of black, white or grey. A few species sport colourful plumes (such as the tropicbirds and some penguins), but most of the colour in seabirds appears in the bills and legs. The plumage of seabirds is thought in many cases to be for camouflage, both defensive (the colour of US Navy battleships is the same as that of Antarctic prions, and in both cases it reduces visibility at sea) and aggressive (the white underside possessed by many seabirds helps hide them from prey below). The usually black wing tips help prevent wear, as they contain melanins to make them black that helps the feathers resist abrasion.

Diet and feeding

Seabirds evolved to exploit different food resources in the world's seas and oceans, and to a great extent, their physiology and behaviour have been shaped by their diet. These evolutionary forces have often caused species in different families and even orders to evolve similar strategies and adaptations to the same problems, leading to remarkable convergent evolution, such as that between auks and penguins. There are four basic feeding strategies, or ecological guilds, for feeding at sea: surface feeding, pursuit diving, plunge-diving, and predation of higher vertebrates; within these guilds, there are multiple variations on the theme.

Surface feeding

Many seabirds feed on the ocean's surface, as the action of marine currents often concentrates food such as krill, forage fish, squid, or other prey items within reach of a dipped head.

Surface feeding itself can be broken up into two different approaches, surface feeding while flying (for example as practiced by gadfly petrels, frigatebirds, and storm petrels), and surface feeding while swimming (examples of which are practiced by fulmars, gulls, many of the shearwaters and gadfly petrels). Surface feeders in flight include some of the most acrobatic of seabirds, which either snatch morsels from the water (as do frigate-birds and some terns), or "walk", pattering and hovering on the water's surface, as some of the storm-petrels do. Many of these do not ever land in the water, and some, such as the frigatebirds, have difficulty getting airborne again should they do so. Another seabird family that does not land while feeding is the skimmer, which has a unique fishing method: flying along the surface with the lower mandible in the water—this shuts automatically when the bill touches something in the water. The skimmer's bill reflects its unusual lifestyle, with the lower mandible uniquely being longer than the upper one.

Surface feeders that swim often have unique bills as well, adapted for their specific prey. Prions have special bills with filters called lamellae to filter out plankton from mouthfuls of water,Brooke, M. (2004). ''Albatrosses and Petrels Across the World''. Oxford University Press, Oxford, UK and many albatrosses and petrels have hooked bills to snatch fast-moving prey. On the other hand, most gulls are versatile and opportunistic feeders who will eat a wide variety of prey, both at sea and on land.

Pursuit diving

Pursuit diving exerts greater pressures (both evolutionary and physiological) on seabirds, but the reward is a greater area in which to feed than is available to surface feeders. Underwater propulsion is provided by wings (as used by penguins, auks, diving petrels and some other species of petrel) or feet (as used by cormorants, grebes, loons and several types of fish-eating ducks). Wing-propelled divers are generally faster than foot-propelled divers. The use of wings or feet for diving has limited their utility in other situations: loons and grebes walk with extreme difficulty (if at all), penguins cannot fly, and auks have sacrificed flight efficiency in favour of diving. For example, the razorbill (an Atlantic auk) requires 64% more energy to fly than a petrel of equivalent size.Gaston, Anthony J.; Jones, Ian L. (1998). ''The Auks'', Oxford University Press, Oxford, Many shearwaters are intermediate between the two, having longer wings than typical wing-propelled divers but heavier wing loadings than the other surface-feeding procellariids, leaving them capable of diving to considerable depths while still being efficient long-distance travellers. The short-tailed shearwater is the deepest diver of the shearwaters, having been recorded diving below .

Some albatross species are also capable of limited diving, with light-mantled sooty albatrosses holding the record at . Of all the wing-propelled pursuit divers, the most efficient in the air are the albatrosses, and they are also the poorest divers. This is the dominant guild in polar and subpolar environments, but it is energetically inefficient in warmer waters. With their poor flying ability, many wing-propelled pursuit divers are more limited in their foraging range than other guilds.

Plunge diving

Gannets, boobies, tropicbirds, some terns, and brown pelicans all engage in plunge diving, taking fast-moving prey by diving into the water from the flight. Plunge diving allows birds to use the energy from the momentum of the dive to combat natural buoyancy (caused by air trapped in plumage), and thus uses less energy than the dedicated pursuit divers, allowing them to utilise more widely distributed food resources, for example, in impoverished tropical seas. In general, this is the most specialised method of hunting employed by seabirds; other non-specialists (such as gulls and skuas) may employ it but do so with less skill and from lower heights. In brown pelicans the skills of plunge-diving take several years to fully develop—once mature, they can dive from 20 m (70 ft) above the water's surface, shifting the body before impact to avoid injury.

It may be that plunge divers are restricted in their hunting grounds to clear waters that afford a view of their prey from the air. While they are the dominant guild in the tropics, the link between plunge diving and water clarity is inconclusive. Some plunge divers (as well as some surface feeders) are dependent on dolphins and tuna to push shoaling fish up towards the surface.

Kleptoparasitism, scavenging and predation

This catch-all category refers to other seabird strategies that involve the next trophic level up. Kleptoparasites are seabirds that make a part of their living stealing food of other seabirds. Most famously, frigatebirds and skuas engage in this behaviour, although gulls, terns and other species will steal food opportunistically. The nocturnal nesting behaviour of some seabirds has been interpreted as arising due to pressure from this aerial piracy. Kleptoparasitism is not thought to play a significant part of the diet of any species, and is instead a supplement to food obtained by hunting. A study of great frigatebirds stealing from masked boobies estimated that the frigatebirds could at most obtain 40% of the food they needed, and on average obtained only 5%. Many species of gull will feed on seabird and sea mammal carrion when the opportunity arises, as will giant petrels. Some species of albatross also engage in scavenging: an analysis of regurgitated squid beaks has shown that many of the squid eaten are too large to have been caught alive, and include mid-water species likely to be beyond the reach of albatrosses. Some species will also feed on other seabirds; for example, gulls, skuas and pelicans will often take eggs, chicks and even small adult seabirds from nesting colonies, while the giant petrels can kill prey up to the size of small penguins and seal pups.

Life history

Seabirds' life histories are dramatically different from those of land birds. In general, they are K-selected, live much longer (anywhere between twenty and sixty years), delay breeding for longer (for up to ten years), and invest more effort into fewer young. Most species will only have one clutch a year, unless they lose the first (with a few exceptions, like the Cassin's auklet), and many species (like the tubenoses and sulids) will only lay one egg a year.

Care of young is protracted, extending for as long as six months, among the longest for birds. For example, once common guillemot chicks fledge, they remain with the male parent for several months at sea. The frigatebirds have the longest period of parental care of any bird except a few raptors and the southern ground hornbill, with each chick fledging after four to six months and continued assistance after that for up to fourteen months. Due to the extended period of care, breeding occurs every two years rather than annually for some species. This life-history strategy has probably evolved both in response to the challenges of living at sea (collecting widely scattered prey items), the frequency of breeding failures due to unfavourable marine conditions, and the relative lack of predation compared to that of land-living birds.

Because of the greater investment in raising the young and because foraging for food may occur far from the nest site, in all seabird species except the phalaropes, both parents participate in caring for the young, and pairs are typically at least seasonally monogamous. Many species, such as gulls, auks and penguins, retain the same mate for several seasons, and many petrel species mate for life. Albatrosses and procellariids, which mate for life, take many years to form a pair bond before they breed, and the albatrosses have an elaborate breeding dance that is part of pair-bond formation.

Breeding and colonies

Ninety-five percent of seabirds are colonial, and seabird colonies are among the largest bird colonies in the world, providing one of Earth's great wildlife spectacles. Colonies of over a million birds have been recorded, both in the tropics (such as Kiritimati in the Pacific) and in the polar latitudes (as in Antarctica). Seabird colonies occur exclusively for the purpose of breeding; non-breeding birds will only collect together outside the breeding season in areas where prey species are densely aggregated.

Seabird colonies are highly variable. Individual nesting sites can be widely spaced, as in an albatross colony, or densely packed as with a murre colony. In most seabird colonies, several different species will nest on the same colony, often exhibiting some niche separation. Seabirds can nest in trees (if any are available), on the ground (with or without nests), on cliffs, in burrows under the ground and in rocky crevices. Competition can be strong both within species and between species, with aggressive species such as sooty terns pushing less dominant species out of the most desirable nesting spaces. The tropical Bonin petrel nests during the winter to avoid competition with the more aggressive wedge-tailed shearwater. When the seasons overlap, the wedge-tailed shearwaters will kill young Bonin petrels in order to use their burrows.

Many seabirds show remarkable site fidelity, returning to the same burrow, nest or site for many years, and they will defend that site from rivals with great vigour. This increases breeding success, provides a place for returning mates to reunite, and reduces the costs of prospecting for a new site. Young adults breeding for the first time usually return to their natal colony, and often nest close to where they hatched. This tendency, known as philopatry, is so strong that a study of Laysan albatrosses found that the average distance between hatching site and the site where a bird established its own territory was ; another study, this time on Cory's shearwaters nesting near Corsica, found that of nine out of 61 male chicks that returned to breed at their natal colony bred in the burrow they were raised in, and two actually bred with their own mother.

Colonies are usually situated on islands, cliffs or headlands, which land mammals have difficulty accessing.Moors, P. J.; Atkinson, I. A. E. (1984). ''Predation on seabirds by introduced animals, and factors affecting its severity''. In ''Status and Conservation of the World's Seabirds''. Cambridge: ICBP. This is thought to provide protection to seabirds, which are often very clumsy on land. Coloniality often arises in types of bird that do not defend feeding territories (such as swifts, which have a very variable prey source); this may be a reason why it arises more frequently in seabirds. There are other possible advantages: colonies may act as information centres, where seabirds returning to the sea to forage can find out where prey is by studying returning individuals of the same species. There are disadvantages to colonial life, particularly the spread of disease. Colonies also attract the attention of predators, principally other birds, and many species attend their colonies nocturnally to avoid predation. Birds from different colonies often forage in different areas to avoid competition.

Migration

Like many birds, seabirds often migrate after the breeding season. Of these, the trip taken by the Arctic tern is the farthest of any bird, crossing the equator in order to spend the Austral summer in Antarctica. Other species also undertake trans-equatorial trips, both from the north to the south, and from south to north. The population of elegant terns, which nest off Baja California, splits after the breeding season with some birds travelling north to the Central Coast of California and some travelling as far south as Peru and Chile to feed in the Humboldt Current. The sooty shearwater undertakes an annual migration cycle that rivals that of the Arctic tern; birds that nest in New Zealand and Chile and spend the northern summer feeding in the North Pacific off Japan, Alaska and California, an annual round trip of .

Other species also migrate shorter distances away from the breeding sites, their distribution at sea determined by the availability of food. If oceanic conditions are unsuitable, seabirds will emigrate to more productive areas, sometimes permanently if the bird is young. After fledging, juvenile birds often disperse further than adults, and to different areas, so are commonly sighted far from a species' normal range. Some species, such as the auks, do not have a concerted migration effort, but drift southwards as the winter approaches. Other species, such as some of the storm petrels, diving petrels and cormorants, never disperse at all, staying near their breeding colonies year round.

Away from the sea

While the definition of seabirds suggests that the birds in question spend their lives on the ocean, many seabird families have many species that spend some or even most of their lives inland away from the sea. Most strikingly, many species breed tens, hundreds or even thousands of miles inland. Some of these species still return to the ocean to feed; for example, the snow petrel, the nests of which have been found inland on the Antarctic mainland, are unlikely to find anything to eat around their breeding sites. The marbled murrelet nests inland in old growth forest, seeking huge conifers with large branches to nest on. Other species, such as the California gull, nest and feed inland on lakes, and then move to the coasts in the winter. Some cormorant, pelican, gull and tern species have individuals that never visit the sea at all, spending their lives on lakes, rivers, swamps and, in the case of some of the gulls, cities and agricultural land. In these cases it is thought that these terrestrial or freshwater birds evolved from marine ancestors. Some seabirds, principally those that nest in tundra, as skuas and phalaropes do, will migrate over land as well.

The more marine species, such as petrels, auks and gannets, are more restricted in their habits, but are occasionally seen inland as vagrants. This most commonly happens to young inexperienced birds, but can happen in great numbers to exhausted adults after large storms, an event known as a ''wreck''.

Relationship with humans

Seabirds and fisheries

Seabirds have had a long association with both fisheries and sailors, and both have drawn benefits and disadvantages from the relationship.

Fishermen have traditionally used seabirds as indicators of both fish shoals, underwater banks that might indicate fish stocks, and of potential landfall. In fact, the known association of seabirds with land was instrumental in allowing the Polynesians to locate tiny landmasses in the Pacific. Seabirds have provided food for fishermen away from home, as well as bait. Famously, tethered cormorants have been used to catch fish directly. Indirectly, fisheries have also benefited from guano from colonies of seabirds acting as fertilizer for the surrounding seas.

Negative effects on fisheries are mostly restricted to raiding by birds on aquaculture, although long-lining

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