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Seabird
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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, while modern seabird families emerged in the Paleogene.
Seabirds generally live longer, breed later and have fewer young than other birds, but they invest a great deal of time in their young. Most species nest in colonies, varying 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 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 oil spills, nets, climate change and severe weather. Conservation efforts include the establishment of wildlife refuges and adjustments to fishing techniques.
Classification
[edit]There exists no single definition of which groups, families and species are seabirds, and most definitions are in some way arbitrary. Elizabeth Shreiber and Joanna 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."[3] However, by convention, all of the Sphenisciformes (penguins), all of the Phaethontiformes (tropicbirds), all of the Procellariiformes (albatrosses and petrels), all of the Suliformes (gannets, boobies, frigatebirds, and cormorants) except the darters, one family of the Pelecaniformes (pelicans), and some of the Charadriiformes (gulls, skuas, 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.[4][5]
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 herons and waders (or shorebirds), such as crab-plovers, are also highly marine, living on the sea's edge (coast), but are also not treated as seabirds. Fish-eating birds of prey, such as sea eagles and ospreys, are also typically excluded, however tied to marine environments they may be.[6] Some birds, such as darters, are primarily found in freshwater habitats, but may occasionally venture into marine or coastal areas as well;[7][8] such birds are generally not considered to be seabirds.
German ornithologist Gerald Mayr defined the "core waterbird" clade Aequornithes in 2010. This lineage gives rise to the Procellariiformes, Sphenisciformes, Suliformes, Pelecaniformes, Ciconiiformes (not seabirds), and Gaviiformes (not seabirds).[9] The tropicbirds (Phaethontiformes) are part of the Eurypygimorphae lineage, which is sister to the Aequornithes;[10] this clade also includes the non-seabird Eurypygiformes (kagu and sunbittern). The Charadriiformes are more distantly related to the other seabirds, being more closely related to the non-seabird Gruiformes (rails and cranes) and Opisthocomiformes (hoatzin) in the clade Gruae.[11]
Evolution and fossil record
[edit]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.[3] They are first known to occur in the Cretaceous period, the earliest being the Hesperornithes. These were flightless seabirds that could dive in a fashion similar to grebes and loons (using its feet to move underwater),[12] but had beaks filled with sharp teeth.[13] Other Cretaceous seabirds included the gull-like Ichthyornithes.[14] Flying Cretaceous seabirds do not exceed wingspans of two meters; piscivorous pterosaurs occupied seagoing niches above this size.[15]

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.[16] 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.[17] 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).[18] 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.[3] Within the Charadriiformes, the gulls and allies (Lari) became seabirds in the late Eocene, and then waders in the middle Miocene (Langhian).[17] The highest diversity of seabirds apparently existed during the Late Miocene and the Pliocene. At the end of the latter, the oceanic food web 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.[19][needs update]
Characteristics
[edit]Adaptations to life at sea
[edit]Seabirds have made numerous adaptations to living on and feeding in the sea. Wing morphology has been shaped by the niche in which 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.[20] 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.[21] 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,[22] and help distinguish familiar nest odours from unfamiliar ones.[23]

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.[25] The excretions from these glands (which are positioned in the head of the birds, emerging from the nasal cavity) are almost pure sodium chloride.[26]
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.[24] 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.[27]
The plumage of most seabirds is less colourful than that of land birds, restricted in the main to variations of black, white or grey.[20] 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,[20] 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 that help the feathers resist abrasion.[28]
Diet and feeding
[edit]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.[29]
Surface feeding
[edit]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 gulls, fulmars, 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 frigatebirds and some terns), or "walk", pattering and hovering on the water's surface, as some of the storm petrels do.[30] 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.[31] 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.[32]
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,[33] 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.[34]
Pursuit diving
[edit]
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.[3] 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.[35] 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 70 metres (230 ft).[36]
Some albatross species are also capable of limited diving, with light-mantled sooty albatrosses holding the record at 12 metres (40 ft).[37] 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.[38]
Plunge diving
[edit]Gannets, boobies, tropicbirds, some terns, and brown pelicans all engage in plunge diving, taking fast-moving prey by diving into the water from 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),[39] 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 (66 ft) above the water's surface, shifting the body before impact to avoid injury.[40]
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.[41] While they are the dominant guild in the tropics, the link between plunge diving and water clarity is inconclusive.[42] Some plunge divers (as well as some surface feeders) are dependent on dolphins and tuna to push shoaling fish up towards the surface.[43]
Kleptoparasitism, scavenging and predation
[edit]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.[44] The nocturnal nesting behaviour of some seabirds has been interpreted as arising due to pressure from this aerial piracy.[45] 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.[3] 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%.[46] 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.[47] 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.[48]
Life history
[edit]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.[3][49] Most species will only have one clutch a year, unless they lose the first (with a few exceptions, like the Cassin's auklet),[50] and many species (like the tubenoses and sulids) will only lay one egg a year.[33]

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.[35] The frigatebirds have the longest period of parental care of any bird except a few raptors and the southern ground hornbill,[51] with each chick fledging after four to six months and continued assistance after that for up to fourteen months.[52] 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.[3]
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.[33] 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.[53]
Breeding and colonies
[edit]
Ninety-five percent of seabirds are colonial,[3] 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.[54]
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.[55] 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.[56]
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.[3] This increases breeding success, provides a place for returning mates to reunite, and reduces the costs of prospecting for a new site.[57] 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 22 metres (72 ft);[58] 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.[59]
Colonies are usually situated on islands, cliffs or headlands, which land mammals have difficulty accessing.[60] This is thought to provide protection to seabirds, which are often very clumsy on land. Coloniality often arises in types of birds 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.[3] 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.[61] Birds from different colonies often forage in different areas to avoid competition.[62]
Migration
[edit]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.[63] 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 64,000 kilometres (40,000 mi).[64]
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.[65] 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.[35] Other species, such as some of the storm petrels, diving petrels and cormorants, never disperse at all, staying near their breeding colonies year round.[66][67][68]
Away from the sea
[edit]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 480 kilometres (300 mi) inland on the Antarctic mainland, are unlikely to find anything to eat around their breeding sites.[69] The marbled murrelet nests inland in old growth forest, seeking huge conifers with large branches to nest on.[70] Other species, such as the California gull, nest and feed inland on lakes, and then move to the coasts in the winter.[71] 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.[20] Some seabirds, principally those that nest in tundra, as skuas and phalaropes do, will migrate over land as well.[4][72]
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.[73]
Relationship with humans
[edit]Seabirds and fisheries
[edit]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,[43] 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.[3] 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.[74]
Negative effects on fisheries are mostly restricted to raiding by birds on aquaculture,[75] although long-lining fisheries also have to deal with bait stealing. There have been claims of prey depletion by seabirds of fishery stocks, and while there is some evidence of this, the effects of seabirds are considered smaller than that of marine mammals and predatory fish (like tuna).[3]

Some seabird species have benefited from fisheries, particularly from discarded fish and offal. These discards compose 30% of the food of seabirds in the North Sea, for example, and compose up to 70% of the total food of some seabird populations.[76] This can have other impacts; for example, the spread of the northern fulmar through the United Kingdom is attributed in part to the availability of discards.[77] Discards generally benefit surface feeders, such as gannets and petrels, to the detriment of pursuit divers like penguins and guillemots, which can get entangled in the nets.[78]
Fisheries also have negative effects on seabirds, and these effects, particularly on the long-lived and slow-breeding albatrosses, are a source of increasing concern to conservationists. The bycatch of seabirds entangled in nets or hooked on fishing lines has had a big impact on seabird numbers; for example, an estimated 100,000 albatrosses are hooked and drown each year on tuna lines set out by long-line fisheries.[79][80][needs update] Overall, many hundreds of thousands of birds are trapped and killed each year, a source of concern for some of the rarest species (for example, only about 2,000 short-tailed albatrosses are known to still exist). Seabirds are also thought to suffer when overfishing occurs.[81] Changes to the marine ecosystems caused by dredging, which alters the biodiversity of the seafloor, can also have a negative impact.[82]
Exploitation
[edit]The hunting of seabirds and the collecting of seabird eggs have contributed to the declines of many species, and the extinction of several, including the great auk and the spectacled cormorant. Seabirds have been hunted for food by coastal peoples throughout history—one of the earliest instances known is in southern Chile, where archaeological excavations in middens has shown hunting of albatrosses, cormorants and shearwaters from 5000 BP.[83] This pressure has led to some species becoming extinct in many places; in particular, at least 20 species of an original 29 no longer breed on Easter Island. In the 19th century, the hunting of seabirds for fat deposits and feathers for the millinery trade reached industrial levels. Muttonbirding (harvesting shearwater chicks) developed as important industries in both New Zealand and Tasmania, and the name of one species, the providence petrel, is derived from its seemingly miraculous arrival on Norfolk Island where it provided a windfall for starving European settlers.[84] In the Falkland Islands, hundreds of thousands of penguins were harvested for their oil each year. Seabird eggs have also long been an important source of food for sailors undertaking long sea voyages, as well as being taken when settlements grow in areas near a colony. Eggers from San Francisco took almost half a million eggs a year from the Farallon Islands in the mid-19th century, a period in the islands' history from which the seabird species are still recovering.[85]
Both hunting and egging continue today, although not at the levels that occurred in the past, and generally in a more controlled manner. For example, the Māori of Stewart Island / Rakiura continue to harvest the chicks of the sooty shearwater as they have done for centuries, using traditional stewardship, kaitiakitanga, to manage the harvest, but now also work with the University of Otago in studying the populations.[86] In Greenland, however, uncontrolled hunting is pushing many species into steep decline.[87]
Other threats
[edit]
Other human factors have led to declines and even extinctions in seabird populations and species. Of these, perhaps the most serious are introduced species. Seabirds, breeding predominantly on small isolated islands, are vulnerable to predators because they have lost many behaviours associated with defence from predators.[60] Feral cats can take seabirds as large as albatrosses, and many introduced rodents, such as the Pacific rat, take eggs hidden in burrows. Introduced goats, cattle, rabbits and other herbivores can create problems, particularly when species need vegetation to protect or shade their young.[88] The disturbance of breeding colonies by humans is often a problem as well—visitors, even well-meaning tourists, can flush brooding adults off a colony, leaving chicks and eggs vulnerable to predators.[89][90]
The build-up of toxins and pollutants in seabirds is also a concern. Seabirds, being apex predators, suffered from the ravages of the insecticide DDT until it was banned; DDT was implicated, for example, in embryo development problems and the skewed sex ratio of western gulls in southern California.[91] Oil spills are also a threat to seabirds: the oil is toxic, and bird feathers become saturated by the oil, causing them to lose their waterproofing.[92] Oil pollution in particular threatens species with restricted ranges or already depressed populations.[93][94]
Climate change mainly affect seabirds via changes to their habitat: various processes in the ocean lead to decreased availability of food and colonies are more often flooded as a consequence of sea level rise and extreme rainfall events. Heat stress from extreme temperatures is an additional threat.[95] Some seabirds have used changing wind patterns to forage further and more efficiently.[96]
In 2023, plasticosis, a new disease caused solely by plastics, was discovered in seabirds. The birds identified as having the disease have scarred digestive tracts from ingesting plastic waste.[97] "When birds ingest small pieces of plastic, they found, it inflames the digestive tract. Over time, the persistent inflammation causes tissues to become scarred and disfigured, affecting digestion, growth and survival."[98]
Conservation
[edit]The threats faced by seabirds have not gone unnoticed by scientists or the conservation movement. As early as 1903, U.S. President Theodore Roosevelt was convinced of the need to declare Pelican Island in Florida a National Wildlife Refuge to protect the bird colonies (including the nesting brown pelicans),[99] and in 1909 he protected the Farallon Islands. Today many important seabird colonies are given some measure of protection, from Heron Island in Australia to Triangle Island in British Columbia.[100][101]
Island restoration techniques, pioneered by New Zealand, enable the removal of exotic invaders from increasingly large islands. Feral cats have been removed from Ascension Island, Arctic foxes from many islands in the Aleutian Islands,[102] and rats from Campbell Island. The removal of these introduced species has led to increases in numbers of species under pressure and even the return of extirpated ones. After the removal of cats from Ascension Island, seabirds began to nest there again for the first time in over a hundred years.[103]
Seabird mortality caused by long-line fisheries can be greatly reduced by techniques such as setting long-line bait at night, dying the bait blue, setting the bait underwater, increasing the amount of weight on lines and by using bird scarers,[104] and their deployment is increasingly required by many national fishing fleets.
One of the Millennium Projects in the UK was the Scottish Seabird Centre, near the important bird sanctuaries on Bass Rock, Fidra and the surrounding islands. The area is home to huge colonies of gannets, puffins, skuas and other seabirds. The centre allows visitors to watch live video from the islands as well as learn about the threats the birds face and how we can protect them, and has helped to significantly raise the profile of seabird conservation in the UK. Seabird tourism can provide income for coastal communities as well as raise the profile of seabird conservation, although it needs to be managed to ensure it does not harm the colonies and nesting birds.[105] For example, the northern royal albatross colony at Taiaroa Head in New Zealand attracts 40,000 visitors a year.[33]
The plight of albatross and large seabirds, as well as other marine creatures, being taken as bycatch by long-line fisheries, has been addressed by a large number of non-governmental organizations (including BirdLife International, the American Bird Conservancy and the Royal Society for the Protection of Birds).[106][107][108] This led to the Agreement on the Conservation of Albatrosses and Petrels, a legally binding treaty designed to protect these threatened species, which has been ratified by thirteen countries as of 2021 (Argentina, Australia, Brazil, Chile, Ecuador, France, New Zealand, Norway, Peru, South Africa, Spain, Uruguay, United Kingdom).[109]
Role in culture
[edit]
Many seabirds are little studied and poorly known because they live far out at sea and breed in isolated colonies. Some seabirds, particularly the albatrosses and gulls, are more well known to humans. The albatross has been described as "the most legendary of birds",[110] and have a variety of myths and legends associated with them. While it is widely considered unlucky to harm them, the notion that sailors believed that is a myth[111] that derives from Samuel Taylor Coleridge's famous poem, "The Rime of the Ancient Mariner", in which a sailor is punished for killing an albatross by having to wear its corpse around his neck. Sailors did, however, consider it unlucky to touch a storm petrel, especially one that landed on the ship.[112]
Gulls are one of the most commonly seen seabirds because they frequent human-made habitats (such as cities and dumps) and often show a fearless nature. Gulls have been used as metaphors, as in Jonathan Livingston Seagull by Richard Bach, or to denote a closeness to the sea; in The Lord of the Rings, they appear in the insignia of Gondor and therefore Númenor (used in the design of the films), and they call Legolas to (and across) the sea. Pelicans have long been associated with mercy and altruism because of an early Christian myth that they split open their breast to feed their starving chicks.[40]
Seabird families
[edit]The following are the groups of birds normally classed as seabirds.[citation needed] For each order, the species counts given are for only the seabird portions (i.e. the listed groups), not the total number of species.
Sphenisciformes (18 species; Antarctic and southern waters)
- Spheniscidae: penguins
Procellariiformes (149 species; pan-oceanic and pelagic)
- Diomedeidae: albatrosses
- Procellariidae: petrels (including fulmars, prions, shearwaters, gadfly petrels, diving petrels, and other petrels)
- Hydrobatidae: northern storm petrels
- Oceanitidae: southern storm petrels
Pelecaniformes (8 species; worldwide)
- Pelecanidae: pelicans
Suliformes (57 species; worldwide)
- Sulidae: gannets and boobies
- Phalacrocoracidae: cormorants
- Fregatidae: frigatebirds
Phaethontiformes (3 species; worldwide tropical seas)
- Phaethontidae: tropicbirds
Charadriiformes (138 species; worldwide)
- Laridae: larids (including gulls, terns, and skimmers)
- Stercorariidae: skuas
- Alcidae: auks
- Genus Phalaropus within Scolopacidae: phalaropes
For an alternative taxonomy of these groups, see also Sibley-Ahlquist taxonomy.
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- ^ Wiedenfeld, D.A. (2016). "Seabird Bycatch Solutions for Fishery Sustainability" (PDF). American Bird Conservancy. Retrieved April 1, 2021.
- ^ Cutlip, Kimbra (August 2, 2017). "Mitigating Seabird Bycatch with Global Fishing Watch". Global Fishing Watch. Retrieved April 1, 2021.
- ^ "Agreement on the Conservation of Albatrosses and Petrels Site". Australian Antarctic Division. Retrieved March 20, 2021.
- ^ Carboneras, C. (1992). "Family Diomedeidae (Albatrosses)". In del Hoyo, J.; Elliott, A.; Sargatal, J. (eds.). Handbook of the Birds of the World. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. pp. 198–215. ISBN 84-87334-10-5.
- ^ Cocker, Mark; Mabey, Richard (2005). Birds Britannica. London: Chatto and Windus. p. 10. ISBN 978-0-7011-6907-7.
- ^ Carboneras, C. (1992). "Family Hydrobatidae (Storm-petrels)". In del Hoyo, J.; Elliott, A.; Sargatal, J. (eds.). Handbook of the Birds of the World. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. pp. 258–271. ISBN 84-87334-10-5.
Further reading
[edit]- Furness, R. W.; P. Monaghan (1987). Seabird Ecology. Tertiary Level Biology. New York: Chapman and Hall. ISBN 978-1-4613-2093-7. OCLC 14069804.
External links
[edit]- Seabirds.net: A data portal for global seabird databases and information outlet for the World Seabird Union
- BirdLife International; Save the Albatross Campaign
- Marine Ornithology, the Journal of Seabird Science and Conservation
Seabird
View on GrokipediaTaxonomy
Definition and Scope
Seabirds are avian species ecologically adapted to exploit marine environments, spending a substantial portion of their lives foraging over open ocean or coastal waters while typically breeding on islands, cliffs, or shorelines.[2] This definition emphasizes their dependence on saltwater habitats for sustenance, with adaptations enabling prolonged time at sea, such as efficient flight or swimming capabilities and physiological mechanisms like supraorbital salt glands for excreting excess sodium.[1] Unlike strictly taxonomic groupings, seabirds form a polyphyletic assemblage defined by functional ecology rather than shared ancestry, uniting diverse lineages that have independently evolved marine lifestyles.[9] The scope encompasses approximately 365 species across at least 17 families, accounting for roughly 3% of global avian diversity, primarily from orders including Procellariiformes (e.g., albatrosses, petrels, and shearwaters), Sphenisciformes (penguins), Suliformes (e.g., gannets, boobies, and cormorants), and select Charadriiformes (e.g., gulls, terns, auks, and skuas).[10] Phaethontidae (tropicbirds), Fregatidae (frigatebirds), and Pelecanidae (pelicans) are also included, though some families like Laridae exhibit partial terrestrial foraging, blurring boundaries with coastal or wetland birds.[11] Exclusions apply to primarily freshwater or inland species, such as certain herons or ducks, even if occasionally marine; the criterion hinges on predominant reliance on oceanic resources for reproduction and survival.[12] This ecological framing highlights seabirds' role as indicators of ocean health, as their distributions and populations reflect prey availability, pollution, and climate shifts, with no single morphological trait universally defining the group beyond habitat affinity.[13]Classification into Families
Seabirds constitute a polyphyletic group, encompassing species from at least five avian orders that have independently adapted to marine lifestyles, rather than forming a single monophyletic clade. This classification reflects ecological convergence rather than shared ancestry, with approximately 363 extant species distributed across 18 families, as recognized by BirdLife International in analyses of global tracking data.[14] Variations in counts arise from differing criteria for "seabird" status, such as the proportion of life spent at sea or reliance on marine prey, leading to estimates ranging from 300 to over 400 species.[10] The primary orders and their constituent seabird families are outlined below, based on modern phylogenetic frameworks like those from the IOC World Bird List, which integrate molecular data to resolve relationships. Penguins (Sphenisciformes) represent a distinct southern-hemisphere radiation, while tube-nosed seabirds (Procellariiformes) dominate pelagic niches. Suliformes and Pelecaniformes include plunge-diving and surface-feeding specialists, and Charadriiformes contribute coastal and pursuit-diving forms. Tropicbirds, sometimes placed in Phaethontiformes, bridge these groups phylogenetically near Procellariiformes.[15]| Order | Family | Representative Genera/Species Count | Key Adaptations/Notes |
|---|---|---|---|
| Sphenisciformes | Spheniscidae | Spheniscus, Aptenodytes (18 spp.) | Flightless swimmers; Antarctic/sub-Antarctic distribution; all species seabirds.[9] |
| Procellariiformes | Diomedeidae | Diomedea (albatrosses, ~21 spp.) | Long-winged gliders; dynamic soaring specialists. |
| Procellariiformes | Procellariidae | Procellaria, Puffinus (petrels/shearwaters, ~100 spp.) | Tube-nosed for salt excretion; diverse foraging strategies. |
| Procellariiformes | Hydrobatidae/Oceanitidae | Hydrobates (storm-petrels, ~25 spp.) | Small, fluttering flyers; oceanic breeders.[15] |
| Phaethontiformes | Phaethontidae | Phaethon (tropicbirds, 3 spp.) | Aerial acrobats; fish-spearing bills; tropical waters. |
| Suliformes | Sulidae | Sula, Morus (gannets/boobies, ~10 spp.) | High-speed plunge divers; colonial nesters. |
| Suliformes | Phalacrocoracidae | Phalacrocorax (cormorants/shags, ~40 spp.) | Pursuit divers; wing-drying behavior post-submersion. |
| Suliformes | Fregatidae | Fregata (frigatebirds, 5 spp.) | Kleptoparasites; inflated throat pouches in males.[9] |
| Pelecaniformes | Pelecanidae | Pelecanus (pelicans, 8 spp.) | Gular pouch for scooping fish; coastal/tropical. |
| Charadriiformes | Laridae | Larus, Sterna (gulls/terns/skimmers, ~100 spp.) | Opportunistic feeders; long migrations. |
| Charadriiformes | Stercorariidae | Stercorarius (skuas/jaegers, 7 spp.) | Predatory/piratical; high-latitude breeders. |
| Charadriiformes | Alcidae | Uria, Fratercula (auks/murres/puffins, ~25 spp.) | Wing-propelled underwater propulsion; northern hemisphere.[9] |
Species Diversity and Endemism
Seabirds encompass approximately 359 species globally, representing about 3.5% of all bird species and spanning multiple orders including Procellariiformes, Sphenisciformes, Pelecaniformes, and select families within Charadriiformes.[6] The Procellariiformes order dominates in species richness, accounting for roughly 149 species such as albatrosses, petrels, and shearwaters, which are adapted for long-distance oceanic foraging.[9] Sphenisciformes contribute 18 species, primarily penguins confined to southern latitudes, while Pelecaniformes include around 57 species across families like Sulidae (gannets and boobies) and Phalacrocoracidae (cormorants). Charadriiformes seabirds, such as auks (Alcidae), gulls (Laridae), and terns (Sternidae), add further diversity through families totaling over 100 species in marine contexts.[9] This distribution reflects evolutionary adaptations to pelagic lifestyles, with higher diversity in temperate and polar regions compared to tropics.[3] Endemism is pronounced among seabirds, driven by the necessity of isolated island breeding colonies that minimize terrestrial predation and provide reliable nesting substrates. Over one-third of procellariiform species, totaling 108, breed exclusively or predominantly on Pacific islands off Mexico, underscoring these archipelagos as critical hotspots.[17] The Benguela Current region off Namibia hosts seven endemic seabird species, including African penguins and Cape gannets, adapted to upwelling-driven productivity.[18] Sub-Antarctic and remote oceanic islands, such as Gough Island in the Tristan da Cunha group, support unique endemics like the Gough bunting (though passerine, seabird-associated ecosystems highlight isolation effects), while the Kermadec Islands harbor the entire breeding population of the endemic Kermadec storm petrel and little shearwater.[19][20] Cabo Verde archipelago features three endemic seabird species and two subspecies, restricted to its volcanic islands.[21] Such patterns arise from philopatry to natal sites and geographic barriers, rendering many species vulnerable to localized threats like invasive predators.[22] Colonial nesting amplifies diversity in hotspots, as seen in dense aggregations of murres and other alcids, where multiple species exploit shared marine resources while partitioning breeding space. Endemic taxa often exhibit restricted ranges, with island-specific radiations in families like Procellariidae, contrasting the wider distributions of cosmopolitan species such as Wilson's storm-petrel. Conservation assessments indicate that endemic seabirds face elevated extinction risks, with 46% of tracked species data revealing breeding concentrations in just 55 countries or territories.[22] Regions like the Southern Ocean south of Tasmania and north-central Atlantic emerge as at-sea foraging hotspots supporting diverse assemblages, though breeding endemism remains tied to land-based isolation.[23]Evolutionary History
Origins in the Cretaceous
The earliest evidence of seabird-like adaptations appears in the Late Cretaceous period, approximately 85 to 66 million years ago, with the evolution of ornithurine birds specialized for marine habitats. Hesperornithiformes, a clade of flightless diving birds including Hesperornis, inhabited the Western Interior Seaway of North America, pursuing fish and other aquatic prey in a manner resembling modern foot-propelled divers.[24] These birds featured elongated bodies, reduced forelimbs, and robust hind limbs with webbed feet for underwater locomotion, marking an early divergence toward fully aquatic lifestyles within Aves.[11] Fossils of Hesperornis regalis, reaching lengths of about 1.8 meters, preserve toothed rostra integrated into the beak structure, facilitating the capture of slippery marine organisms amid competition from reptilian predators like mosasaurs.[25] Contemporaneous ichthyornithiforms, such as Ichthyornis dispar, represented volant counterparts with keeled sterna supporting flight and similarly dentulous jaws, suggesting a spectrum of aerial and diving strategies in proto-seabird lineages.[26] These forms, preserved in lagerstätten like the Niobrara Formation, indicate that selective pressures from expanding epicontinental seas drove initial morphological innovations for pelagic foraging, predating the diversification of toothless neornithine seabirds.[11] This Cretaceous radiation occurred within Ornithurae, the broader group encompassing modern birds, but hesperornithiforms and ichthyornithiforms did not survive the end-Cretaceous mass extinction, yielding to post-K-Pg adaptive expansions among surviving avian clades.[27] Their existence underscores a pre-extinction experimentation with marine niches, supported by biomechanical adaptations evident in skeletal remains, though limited global distribution reflects the era's fragmented ocean connectivity.[24]Fossil Evidence and Key Transitions
The fossil record of seabirds originates in the Late Cretaceous, with evidence of early aquatic adaptations among ornithurine birds predating the K-Pg extinction by millions of years. Key specimens include those of Hesperornithiformes and Ichthyornithiformes, which display a mosaic of primitive and derived traits indicative of transitions toward specialized marine foraging. These fossils, recovered from marine deposits such as the Western Interior Seaway, reveal the development of diving propulsion via hind-limb modifications and cranial features for grasping prey, distinct from terrestrial avian ancestors.[28] Hesperornis regalis, dating to approximately 83.6–72 million years ago, exemplifies flightless specialization, featuring a dentulous bill integrated into a robust skull, reduced wings, and powerful, paddle-like feet for underwater pursuit of fish and ammonites. Analysis of bone microstructure indicates rapid skeletal maturation within one year, supporting high metabolic rates akin to those enabling endurance in modern diving birds. This lineage's secondary loss of flight underscores a causal trade-off favoring aquatic efficiency over aerial mobility, a pattern echoed in later seabird groups.[28][29][30] Ichthyornis dispar, from around 85 million years ago, represents a flying counterpart with transitional cranial morphology: teeth set within an incipient keratinous beak, bridging theropod dentition and the edentulous rhamphotheca of crown-group birds. Micro-CT reconstructions of its skull highlight expanded braincase volume for enhanced sensory processing, alongside a lightweight skeleton suited for agile flight over water. These features facilitated a gull-like ecology, capturing evasive marine prey, and mark a critical step in the evolution of beak diversification for varied trophic roles.[31][32] Together, these taxa provide empirical evidence of pre-extinction experimentation with marine lifestyles, including dentition suited for slippery quarry and locomotor shifts prioritizing submersion over sustained flight. Such innovations, preserved in lagerstätten like the Smoky Hill Chalk, inform causal mechanisms driving seabird diversification, where environmental pressures from epicontinental seas selected for physiological tolerances to salinity and hypoxia.[28]Adaptive Radiations Post-Mass Extinctions
The Cretaceous-Paleogene (K-Pg) mass extinction event, dated to 66 million years ago, eliminated non-avian dinosaurs, pterosaurs, marine reptiles including mosasaurs and plesiosaurs, and numerous archaic bird lineages such as enantiornithines, thereby vacating extensive pelagic and coastal niches. This ecological release enabled surviving crown-group birds (Neornithes) to undergo adaptive radiations into marine habitats, with fossil records indicating a swift diversification of seabird morphologies tailored to oceanic foraging. The extinction's selective pressures favored ground- and water-associated birds over arboreal forms, setting the stage for neornithine seabirds to exploit abundant post-extinction marine resources like fish stocks recovering from the collapse of reptilian predators.[33][27][34] Among the earliest post-K-Pg seabird fossils is a diminutive pelagornithid specimen from the early Paleocene of New Zealand, approximately 60-61 million years old, which documents a basal member of this group and suggests an origin in the Southern Hemisphere amid rapid neornithine expansion into open-ocean ecosystems altered by the extinction. Pelagornithidae, featuring elongated bills with bony pseudoteeth for grasping elusive prey and wingspans reaching 6 meters for efficient soaring, proliferated globally from the Paleocene through the Miocene, embodying key innovations in sustained flight and surface-piercing predation that filled niches left by vanished flying reptiles and toothed seabirds. Their near-immediate appearance underscores the opportunistic radiation driven by reduced competition and enhanced prey availability in warming Paleogene seas.[35] By the Eocene, around 50 million years ago, further radiations encompassed early procellariiforms (such as petrels and albatrosses) and sphenisciforms (penguins), alongside extinct plotopterids—penguin-like wing-propelled divers from the Eocene-Oligocene—that specialized in underwater pursuit of fish and squid. These developments aligned with the Paleocene-Eocene Thermal Maximum's global warming, which boosted ocean productivity and facilitated niche partitioning among flighted soarers, plunge-divers, and pursuit divers. Unlike the pre-extinction Cretaceous seabird assemblage dominated by toothed hesperornithiforms, post-K-Pg forms emphasized keratinous bills and varied propulsion strategies, reflecting causal adaptations to a predator-scarce marine realm and establishing the foundational diversity of modern seabird orders.[36][37]Morphology and Physiology
Structural Adaptations for Marine Life
Seabirds exhibit dense, interlocking feather structures that interlock via barbules to form a barrier against water penetration, supplemented by oils from the uropygial gland applied during preening to maintain waterproofing during prolonged marine exposure.[38] This adaptation minimizes heat loss and prevents plumage from becoming waterlogged, essential for species foraging far offshore.[39] Supraorbital salt glands, positioned above the eyes and connected to nasal passages, enable seabirds to excrete excess sodium chloride in a concentrated solution hypertonic to seawater, countering osmotic stress from drinking saline water and consuming salty prey.[40] These glands, derived from lateral nasal glands, activate via neural and hormonal signals in response to salt loads, allowing survival without frequent freshwater access.[41] Webbed feet with totipalmate or semipalmate configurations provide propulsion for swimming, varying by foraging depth: surface swimmers like petrels have partial webbing, while pursuit divers like cormorants possess fully webbed feet for efficient underwater paddling.[42] Streamlined fusiform body shapes reduce drag during dives and surface travel, with many species featuring short necks and tails for hydrodynamic efficiency.[43] Wing morphology diversifies by lifestyle: long, narrow wings in albatrosses and shearwaters facilitate dynamic soaring over vast ocean expanses with minimal energy expenditure, while short, stiffened wings in auks and penguins function as flippers for underwater propulsion, often paired with reduced pneumatization of bones to increase overall density and aid submersion.[11] Bills are typically hooked or pointed for grasping slippery fish, with pelicans featuring expandable pouches and gannets tapered for plunge-diving precision.[12] These skeletal and integumentary features collectively support the dual demands of aerial and aquatic locomotion inherent to marine existence.[39]Sensory and Physiological Specializations
Seabirds possess acute visual capabilities tailored for detecting prey across expansive marine vistas, with many species exhibiting high spatial resolution and enhanced optical sensitivity to low light conditions during dawn or dusk foraging. Procellariiform seabirds, such as albatrosses and petrels, demonstrate particularly refined vision adapted to dynamic ocean surfaces, allowing precise targeting of shoaling fish or plankton blooms from altitudes exceeding 10 meters.[44] Olfaction plays a prominent role in prey location for procellariiforms, which feature enlarged olfactory bulbs relative to other birds, enabling detection of volatile compounds like dimethylsulfide (DMS) emitted by phytoplankton and krill aggregations. Wandering albatrosses (Diomedea exulans), for example, respond to fishy odors in field trials, using smell to home in on productive patches over hundreds of kilometers.[45] [46] This sensory reliance contrasts with diurnal visual foragers like gulls, underscoring olfactory evolution tied to nocturnally active or pelagic lifestyles.[47] Auditory sensitivity in seabirds centers on frequencies of 1.0–3.0 kHz for intraspecific communication, territorial defense, and predator evasion, with diving taxa showing impedance-matching adaptations for underwater sound propagation despite reduced aerial efficiency when submerged.[48] Tactile mechanoreceptors in bills, concentrated in species like shearwaters and penguins, detect substrate vibrations or prey movements during tactile foraging in turbid waters or at night, enhancing localization where vision fails.[49] Physiologically, seabirds maintain ionic balance through supraorbital salt glands that secrete hypertonic NaCl solutions—up to twice seawater concentration—functioning as auxiliary kidneys to counter salt loads from ingested seawater and marine prey. These glands, innervated by parasympathetic pathways, activate rapidly in response to hyperosmotic stress, excreting 4–5% of body weight in saline daily in species like herring gulls (Larus argentatus).[50] [51] Diving seabirds, including auks and penguins, exhibit elevated myoglobin concentrations in flight muscles—up to 10 times terrestrial avian levels—for extended aerobic dives, supplemented by peripheral vasoconstriction and cardiac shunts that prioritize cerebral and myocardial oxygenation while minimizing nitrogen narcosis risks at depths beyond 100 meters.[52] These adaptations, coupled with denser bone marrow in pursuit divers, facilitate breath-hold durations of 2–5 minutes, balancing energetic costs of repeated immersion against aerial efficiency demands.[53]Variations Across Seabird Groups
Seabirds display substantial morphological and physiological diversity reflecting adaptations to distinct marine foraging strategies, from aerial pursuit to deep-water diving. Body sizes range from the 40-gram Wilson's storm-petrel (Oceanites oceanicus), a small Procellariiform reliant on surface prey, to the 12-kilogram wandering albatross (Diomedea exulans), optimized for long-distance gliding over open oceans.[1] Plumage across groups is typically dichromatic in black, white, and gray tones for countershading and camouflage, with denser, scale-like feathers in diving specialists to enhance insulation and waterproofing.[54] In Sphenisciformes (penguins), flightlessness is universal, with wings modified into rigid, flipper-like structures via fused bones for underwater propulsion, enabling pursuits of fish and krill at depths exceeding 500 meters in species like the emperor penguin (Aptenodytes forsteri). Legs are positioned posteriorly for steering, and non-pneumatized bones reduce buoyancy, paired with elevated myoglobin levels in muscles for prolonged aerobic dives. These traits contrast sharply with volant groups, emphasizing energy allocation to swimming over aerial locomotion.[54] Procellariiformes (albatrosses, petrels, shearwaters) feature tubular nostrils aiding olfaction for locating prey, with supraorbital salt glands excreting concentrated brine to manage osmotic stress from marine diets. Wing morphology varies: albatrosses possess high-aspect-ratio wings with low loading for dynamic soaring in windy regimes, while diving petrels like Pelecanoides urinatrix have shorter, stubbier wings for paddling. Dive capabilities differ markedly within the order; sooty shearwaters (Puffinus griseus) achieve deeper (up to 70 meters) and longer dives with higher hematocrit and red blood cell counts for enhanced oxygen transport, compared to shallower, more frequent dives by common diving petrels relying on greater respiratory oxygen stores. Wing loading correlates positively with median wind speeds at breeding sites, allowing tolerance of gales up to 50 meters per second in polar species.[54][55][56] Suliformes (cormorants, gannets, boobies) exhibit streamlined bodies for underwater agility, with totipalmate feet fully webbed for propulsion and bills adapted for spearing: gannets (Morus spp.) have hinged crania to withstand plunge-dive impacts from heights of 30 meters, reaching speeds over 100 kilometers per hour. Cormorants chase prey subaquatically with partially wettable plumage to reduce drag, contrasting the fully preened waterproofing in surface feeders. Salt glands are prominently orbital, processing high-salinity loads efficiently.[54] Among Charadriiformes (alcids, gulls, terns), alcids like murres (Uria spp.) converge on penguin-like diving via compact torsos, short wings for wing-beat propulsion to 200 meters, and dense bones for ballast, forgoing the pneumatic skeletons of aerial specialists. Gulls and terns, conversely, employ agile, flapping flight with forked tails and pointed bills for surface skimming or hovering over fish schools, with less emphasis on diving physiology and more on visual acuity for opportunistic foraging. These variations underscore niche partitioning, where pursuit divers prioritize oxygen storage and skeletal density, while gliders emphasize aerodynamic efficiency.[54]Foraging Ecology
Dietary Preferences and Trophic Levels
Seabirds primarily consume marine prey including fish, cephalopods, and crustaceans, with dietary composition varying by species, foraging habitat, and environmental conditions.[57] Analysis of regurgitated boluses and stomach contents from procellariiform seabirds, such as petrels and albatrosses, frequently identifies epipelagic fish and squid as dominant components, often comprising over 50% of identifiable prey items in breeding colonies.[58] Crustaceans, including euphausiids like krill, constitute a major proportion in the diets of penguins and certain alcids, with studies reporting up to 90% krill in Adélie penguin diets during austral summer. Scavenging on fishery discards or offal supplements diets for opportunistic species like gulls and shearwaters, though this varies regionally and with fishing intensity.[59] Stable isotope analysis using δ¹⁵N signatures positions most seabirds at trophic levels of 3 to 4 within pelagic food webs, reflecting their role as predators of secondary consumers such as small fish and squid that feed on zooplankton.[60] Plankton- or crustacean-dependent species, including some storm-petrels, exhibit lower trophic positions around 3.4–3.5, while piscivores like shags and cormorants reach 3.7–3.9, indicating greater reliance on higher-order prey.[61] Comparisons across taxa confirm that δ¹⁵N-derived trophic inferences align with conventional dietary assessments, though isotopes integrate long-term assimilation and may reveal subtler shifts undetectable in snapshot prey samples.[62] Long-term monitoring in northern hemisphere populations has documented declines in mean trophic position for species like black-legged kittiwakes, from approximately 3.8 to 3.5 between 1978 and 2015, correlating with reduced availability of lipid-rich, high-trophic prey amid ocean warming.[63] Dietary guilds among seabirds include specialists on small planktonic organisms, generalists targeting schooling fish, predators of large nekton like squid, and scavengers exploiting anthropogenic food sources, influencing their vulnerability to prey fluctuations.[59] For instance, DNA metabarcoding of buccal swabs from Manx shearwaters identifies fish as the most frequent prey category (over 60% occurrence), followed by cephalopods, underscoring molecular methods' utility in resolving fine-scale trophic interactions.[64] These preferences underscore seabirds' position as mid-to-upper trophic regulators, exerting top-down pressure on forage fish stocks estimated at 10–50 million metric tons annually across global populations.[65]Hunting Techniques and Strategies
Seabirds employ a diverse array of hunting techniques adapted to the challenges of capturing prey in marine environments, ranging from surface waters to depths exceeding 100 meters. These strategies include surface seizing, plunge diving, pursuit diving, and kleptoparasitism, often tailored to specific prey types such as fish, squid, and plankton.[66] Foraging success depends on morphological adaptations, sensory cues like olfaction and vision, and behavioral plasticity, with many species exhibiting individual specialization in techniques.[67] Surface seizing predominates among procellariiforms like storm-petrels and shearwaters, where birds flutter low over waves to peck plankton, krill, or small fish directly from the water column without submerging.[68] Storm-petrels patter their feet on the surface to agitate and capture zooplankton, leveraging erratic flight to exploit concentrated patches formed by ocean currents.[69] Gulls and some terns similarly seize prey from the surface, scavenging or targeting opportunistically available forage fish and squid.[11] Plunge diving is characteristic of sulids such as gannets and boobies, who spot prey from heights of 10-40 meters and dive vertically at speeds over 80 km/h, using streamlined bodies and air sacs to cushion impact and pursue fish underwater briefly.[70] [71] Brown pelicans execute high-speed plunges resembling split-S maneuvers, folding wings mid-dive to strike fish schools with precision, while terns perform shallower versions for aerial spotting and rapid entry.[72] These techniques minimize injury through skeletal reinforcements and flexible necks, enabling repeated dives during foraging bouts.[73] Pursuit diving relies on underwater propulsion, primarily by wing-beating in alcids (auks) and penguins, who chase schooling fish like capelin or herring to depths of 100-200 meters in species such as murres.[74] Auks flap wings efficiently in water for "flight-like" pursuit, contrasting higher energetic costs in air, which constrains their foraging range.[75] Penguins similarly herd and corral prey using coordinated group dives, facilitating capture of evasive fish.[76] Kleptoparasitism serves as a low-risk strategy for skuas and frigatebirds, who harass other seabirds mid-flight to induce regurgitation of captured prey, often targeting piscivores like terns or gannets.[77] Skuas pursue victims persistently, while frigatebirds use agile soaring to intercept, supplementing direct predation during breeding seasons when energy demands peak.[78] This behavior exploits the foraging efforts of conspecifics or sympatric species, enhancing efficiency in unpredictable prey distributions.[79] Many seabirds integrate social strategies, foraging in multispecies flocks to cue on predator activity like tuna schools driving prey to the surface, amplifying individual detection via visual or olfactory signals.[80] [81] Such associative foraging reduces search costs but varies by taxonomy and resource patchiness.[82]Interactions with Prey Populations
Seabirds, as central-place foragers during breeding seasons, create localized zones of prey depletion surrounding their colonies, a phenomenon known as Ashmole's halo, where intensified predation reduces prey densities in proximity to nesting sites.[83] This effect arises from the constraint that breeding seabirds must return to colonies to provision chicks, concentrating foraging effort within accessible radii and leading to measurable reductions in prey biomass; for instance, masked boobies (Sula dactylatra) at Ascension Island depleted flying fish (Exocoetidae) populations by up to 50% within 10-20 km of the colony compared to distant areas, as evidenced by acoustic surveys and dietary analyses conducted in 2019-2020.[83] Such depletion supports the hypothesis that food limitation regulates seabird population sizes, with higher-density colonies exhibiting stronger halo effects due to cumulative foraging pressure.[84] Beyond immediate depletion, seabird predation influences prey population dynamics through selective foraging on abundant or vulnerable schools, often targeting juvenile or schooling fish species like anchovies (Engraulis spp.) and sardines (Sardinops spp.), which can alter prey age structures and recruitment rates in coastal ecosystems.[85] Studies in the California Current system demonstrate that Brandt's cormorants (Uria lugge) switch prey in response to environmental variability, consuming more juvenile salmon (Oncorhynchus spp.) during low anchovy availability, thereby imposing variable predation mortality that correlates with oceanographic conditions like upwelling intensity.[86] In the Southern Ocean, Adélie penguins (Pygoscelis adeliae) harvest Antarctic krill (Euphausia superba) at rates reflecting broader prey pulses, but their impact remains subordinate to abiotic factors and large whales, with annual consumption estimates around 100-200 million tons across all krill predators insufficient to drive basin-scale declines absent other stressors.[87] Prey populations exhibit adaptive responses to seabird foraging, including behavioral shifts such as deeper diving or dispersion to evade surface predators like shearwaters and petrels, which in turn can feedback to limit seabird breeding success when prey evades capture.[88] Empirical models indicate that density-dependent competition among seabirds amplifies these interactions, with larger colonies forcing individuals to forage farther and encounter lower per capita prey encounter rates, stabilizing predator-prey oscillations through enhanced predation on denser prey patches.[89] While global seabird predation rarely causes widespread prey crashes—due to the mobility of marine prey and seabirds' opportunistic diets—localized effects around island colonies can persist for months post-breeding, influencing nutrient cycling via guano deposition but without evidence of long-term trophic cascades in most systems.[90]Reproduction and Demography
Breeding Systems and Parental Care
Seabirds primarily exhibit social monogamy, forming long-term pair bonds that are renewed annually at breeding colonies, with divorce rates remaining low under stable conditions but rising after reproductive failures or environmental stressors like warming ocean temperatures that impair foraging.[91][92] Long-term partners display reduced courtship intensity and more equitable sharing of duties compared to newly formed pairs, minimizing sexual conflict over care allocation.[93] Genetic studies reveal occasional extra-pair paternity, yet overall pair fidelity supports biparental investment in a single breeding attempt per season.[94] Breeding is highly colonial, with over 95% of species aggregating in dense groups on predator-poor islands or cliffs, where benefits such as diluted predation risk outweigh costs like conspecific aggression.[95] Courtship involves species-specific displays, including mutual ornamentation assessments in crested auklets and synchronized vocalizations or dances in albatrosses, facilitating mate choice and bond reinforcement.[96] Nests vary by taxon: burrows or crevices for petrels and shearwaters, exposed ledges for murres, or stick platforms for boobies and pelicans. Clutch sizes generally range from one egg in procellariiforms like albatrosses and petrels to two or three in alcids, gulls, and terns, reflecting trade-offs between offspring number and per-chick investment.[97] Incubation periods span 30 to 80 days, with biparental reliefs ensuring continuous coverage; coordination peaks during this phase, as partners alternate shifts to forage.[98] Established pairs achieve more balanced incubation, producing larger eggs than less compatible newcomers.[93] Chick-rearing demands sustained biparental provisioning, with parents undertaking extended foraging bouts to deliver energy-rich marine prey, often coordinating departures and arrivals to maintain feeding rates and support chick growth.[99] This coordination diminishes as chicks develop independence, allowing greater parental flexibility amid variable prey availability.[98] Seabirds adopt a conservative strategy, curtailing effort under poor conditions to preserve adult condition for future breeding, given their longevity exceeding 20-50 years in many species.[100] Variations occur across families; for instance, thick-billed murres feature one parent shadowing the fledgling to sea, while little auks adjust dive depths and trip durations flexibly to match environmental demands.[101][102]Colony Dynamics and Site Fidelity
Seabirds predominantly breed in colonies, with approximately 95% of species utilizing synchronous aggregations at limited sites, which facilitates predator dilution and communal defense against threats such as aerial and terrestrial predators.[103] Colony size influences reproductive outcomes, as larger colonies often exhibit higher and more stable breeding success due to social facilitation and reduced per capita predation risk, though subcolony variations in environmental conditions like sea surface temperature can lead to disparities in foraging efficiency and chick fledging rates—for instance, one subcolony in a study of little penguins fledged 30% more chicks than another owing to cooler waters supporting better prey availability.[104] [105] However, dense colonies can incur costs from conspecific aggression, increased disease transmission, and intensified competition for nest sites and food, potentially undermining benefits in overcrowded conditions.[106] Colony dynamics are shaped by metapopulation processes, including immigration, emigration, and local extinction risks, with climate variability driving shifts in occupancy and size; for example, northern gannet colonies respond to warming oceans through altered foraging and dispersal patterns.[107] Breeding synchrony within colonies enhances collective vigilance and information transfer about foraging opportunities, allowing less experienced individuals to follow successful foragers, though this advantage diminishes if prey patches become unpredictable.[108] In recovering populations, such as common murres, colony growth correlates with elevated breeding success, reaching up to 190 pairs by 2004 in re-established sites, underscoring density-dependent positive feedbacks.[109] Adult seabirds demonstrate variable but often substantial site fidelity to breeding colonies, with philopatry rates lower overall than previously assumed and exhibiting wide interspecies differences, influenced by prior reproductive performance and environmental cues.[110] In Monteiro's storm-petrel, fidelity to specific nests strengthens following successful breeding, particularly under unfavorable oceanic conditions like low chlorophyll-a concentrations, enabling birds to prioritize high-quality sites that predict future success (β = 4.16 for success effect).[111] This behavior promotes population stability by retaining experienced breeders but can trap individuals in declining habitats, limiting adaptive dispersal; for instance, northern gannets maintain strong colony fidelity even after high-mortality events, constraining metapopulation recovery.[112] Failed breeders and immatures show reduced fidelity compared to successful adults, reflecting conditional strategies balancing familiarity benefits against prospecting for alternatives.[113]Life History Trade-offs and Longevity
Seabirds exemplify slow life-history strategies, prioritizing high adult survival and longevity over rapid reproduction, adaptations honed by the unpredictable availability of marine resources. Adult annual survival rates frequently exceed 0.90 in long-lived groups like Procellariiformes, enabling maximum lifespans well over 50 years; for instance, Laysan albatrosses (Phoebastria immutabilis) have documented lifespans surpassing 60 years in the wild, with some individuals reaching beyond 70.[114][115] This extended lifespan supports delayed maturity—often 5–10 years or more—and low annual fecundity, typically one chick per breeding attempt, allowing cumulative reproductive output across multiple seasons despite high chick mortality risks.[116] A core trade-off underlies this strategy: investment in current reproduction compromises future survival and subsequent breeding probability. Comparative analyses across 44 species of albatrosses and petrels reveal a significant negative correlation between annual reproductive output (e.g., chick production and fledging success) and adult survival, persisting after phylogenetic and body-size corrections, with reproductive effort also trading off against age at maturity.[117] In species like the northern fulmar (Fulmarus glacialis), experimental nest failures demonstrate that skipping breeding enhances long-term survival and return rates, while successful breeding elevates mortality risks, particularly for females due to asymmetric costs in foraging and incubation.[118][119] These costs extend to senescence, where early-life breeding accelerates reproductive decline, as observed in long-finned pilot whales and black-legged kittiwakes (Rissa tridactyla), with trade-offs evident independent of seasonal breeding timing.[120] Longevity thus buffers these trade-offs, concentrating lifetime reproductive success in later years; in kittiwakes, 80–83% of total output derives from extended adult survival rather than early fecundity.[114] Environmental variability amplifies such dynamics, with individuals in resource-poor years often deferring or skipping breeding to preserve condition, a tactic supported by high baseline survival that sustains population stability over decades.[121] This K-selected approach contrasts with faster-paced terrestrial birds, reflecting causal pressures from marine habitat patchiness and high juvenile mortality, which favor survival maximization in adults.[114]Movement and Distribution
Migration Patterns and Navigational Mechanisms
Many seabirds, particularly species in the orders Procellariiformes and Charadriiformes, undertake long-distance migrations between breeding colonies—often located in high-latitude or temperate regions—and non-breeding grounds in subtropical or tropical oceans, driven by seasonal prey availability and breeding phenology.[122] These patterns vary by taxon; for instance, albatrosses and petrels frequently exhibit circumpolar or trans-oceanic routes, while some gulls and terns perform partial migrations or remain resident in productive coastal zones.[123] Tracking studies using geolocators and satellite tags reveal that migratory flights often involve increased daily flight distances and durations compared to breeding periods, with birds allocating more time to soaring and gliding to optimize energy expenditure over vast pelagic expanses.[123] The Arctic tern (Sterna paradisaea) exemplifies extreme migratory commitment, breeding in Arctic and subarctic regions before traveling southward to Antarctic waters, achieving the longest annual migration documented in any animal at approximately 70,000–96,000 km round-trip.[124] Geolocator data from tracked individuals confirm this pole-to-pole circuit, with juveniles following similar routes to adults after an initial orientation phase, ensuring access to perpetual daylight and high-productivity foraging zones year-round.[124] Similarly, the sooty shearwater (Ardenna grisea) departs breeding colonies in New Zealand and southern Chile to traverse the Pacific in a figure-eight pattern, covering over 65,000 km to exploit northern hemisphere upwellings, as evidenced by archival tag deployments on 19 birds that mapped resource integration across hemispheres.[125] Seabirds navigate these routes using a multimodal system integrating geomagnetic, celestial, and olfactory cues, with evidence from displacement experiments and sensory manipulations indicating redundancy to compensate for environmental variability.[126] Procellariiform seabirds, such as shearwaters, imprint on the Earth's magnetic field parameters (inclination and intensity) during fledging for initial orientation, as demonstrated in Manx shearwater (Puffinus puffinus) fledglings that recalibrated to natal sites after magnetic relocation.[126] Olfactory mechanisms play a key role in homing for petrels and albatrosses, where anosmic birds (with temporarily blocked olfactory nerves) fail to return from short-range displacements over open ocean, underscoring smell-based mapping of wind-borne odor plumes from productive waters.[127] Celestial compasses, including sun arc and polarized light patterns, provide time-compensated orientation during diurnal flights, while stellar cues may assist nocturnal migrants, though empirical validation remains stronger for geomagnetic and olfactory modalities in seabirds.[127]Range Expansions and Contractions
Seabird ranges have shifted in response to environmental drivers, including ocean warming, prey redistribution, and habitat modifications, with patterns varying by species and region. Poleward expansions at leading range edges often occur as warming oceans displace prey toward higher latitudes, enabling colonization of novel breeding and foraging areas, though trailing edge contractions frequently result in net range reductions. For instance, analyses of marine species distributions indicate abundance increases at poleward boundaries linked to thermal tolerance limits, while equatorward declines reflect unsuitable conditions.[128][129] Procellariiform seabirds, such as albatrosses, petrels, shearwaters, and storm petrels, demonstrate range contractions amid rapid climate change, with shrinking habitable areas elevating extinction risks through reduced population connectivity and dispersal limitations. Projections for Southern Ocean albatrosses and petrels consistently forecast poleward distributional shifts under multiple climate scenarios, yet overall range sizes contract due to habitat compression at equatorial trailing edges outpacing gains elsewhere. In the North Pacific, Laysan albatrosses (Phoebastria immutabilis) successfully expanded their breeding range northward, establishing new colonies while adapting foraging behaviors to exploit altered prey availability.[130][131][132] Contractions arise from habitat loss, including sea-level rise eroding low-lying breeding islands critical for burrow- and surface-nesting species, and intensified storms disrupting colony persistence. Multi-decadal surveys of Arctic-associated seabirds like little auks and Brünnich’s guillemots reveal distribution shifts tied to climate impacts on prey, with abundance declines in core ranges despite some poleward movements. Genomic studies of southern seabird species further suggest adaptive introgression facilitates responses to range alterations, but persistent contractions threaten endemic populations in warming hotspots.[133][134][135]Responses to Environmental Variability
Seabirds demonstrate behavioral plasticity in their movement patterns to mitigate short-term environmental fluctuations, such as those induced by the El Niño-Southern Oscillation (ENSO), which alters ocean temperatures, upwelling, and prey distribution. During intense El Niño events, tropical seabirds exhibit heightened sensitivity to precursors like anomalous sea surface temperatures months before peak warming, prompting shifts in foraging ranges and reduced breeding participation to track ephemeral prey patches. For instance, in the southeastern Pacific, El Niño reduces the anchovy food base for guano birds like Peruvian pelicans and boobies, leading to widespread nest desertion and extralimital dispersal as individuals relocate to areas with persistent productivity.[136][137][138] Wind regime changes during ENSO events further influence pelagic seabird distribution, with species like Laysan albatrosses (Phoebastria immutabilis) experiencing elevated wind speeds that enhance flight efficiency but disrupt incubation schedules on breeding grounds in the North Pacific. Black-footed albatrosses (P. nigripes), foraging more southerly, show muted responses, highlighting species-specific adaptations tied to baseline habitat overlap with variability hotspots. In the Southern Ocean, Antarctic seabirds such as Adélie penguins (Pygoscelis adeliae) and thin-billed prions (Pachyptila belcheri) display contrasting long-term demographic responses to sea ice and temperature variability over 40-year records, with penguins advancing breeding phenology to exploit early ice melt while prions suffer deferred recruitment amid reduced krill availability.[139][140] Individual-level flexibility enables some seabirds to adjust migration routes dynamically to oceanographic shifts, as evidenced by GPS tracking of Manx shearwaters (Puffinus puffinus), where birds shortened non-breeding sojourns in cooler North Atlantic waters during warmer summers, correlating with sea surface temperature anomalies exceeding 1°C above averages in 2014–2020. Such plasticity buffers against variability but varies by life stage; juveniles often explore broader ranges during poor conditions, while adults prioritize fidelity to productive corridors. However, mechanistic models underscore limits, as sustained wind-driven energetic costs during prolonged anomalies can constrain range expansions, particularly for central-place foragers during breeding.[141][142][123] Breeding range shifts represent a distributional response to decadal variability, with tropical species like brown boobies (Sula leucogaster) and blue-footed boobies (S. nebouxii) colonizing higher-latitude sites such as Sutil Island off Mexico by 2024, tracking poleward prey migrations amid 0.5–1.0°C regional warming since 1980. These expansions contrast with contractions in temperate populations facing intensified storm frequency, where storm-petrels (Oceanodroma spp.) alter diel flight budgets to evade turbulence, increasing energy expenditure by up to 20% during migration. Empirical tracking data reveal that while short-term variability elicits reversible behavioral tweaks, cumulative effects from recurrent events like ENSO amplify risks of maladaptation in less plastic species.[143][144][122]Ecological Roles
Indicators of Marine Ecosystem Health
Seabirds serve as effective bioindicators of marine ecosystem health owing to their positions as upper-trophic-level predators that integrate signals from large foraging areas, bioaccumulate contaminants through diet, and exhibit measurable responses in population dynamics and reproductive output to changes in prey abundance and environmental conditions.[145] Their breeding colonies, often monitored long-term, provide data on food web stability, as declines in breeding success correlate with reduced forage fish stocks influenced by overfishing or oceanographic shifts.[146] For instance, global analyses of monitored seabird populations reveal an overall decline of 69.7% from 1950 to 2010, equivalent to a loss of approximately 230 million individuals, signaling broad degradation in marine productivity.[147][148] Reproductive metrics, such as fledging rates and chick survival, reflect prey availability and ocean health; in regions like the Southern Ocean, hemispheric asymmetries in warming have led to divergent breeding successes, with northern hemisphere populations faring worse amid intensified human impacts and temperature anomalies.[149] Seabird foraging behaviors, tracked via biologging, further indicate ecosystem shifts, as extended trip durations or reduced meal sizes during breeding seasons align with diminished prey densities from climate-driven habitat alterations.[150] These responses underscore seabirds' utility in detecting trophic cascades, where forage fish depletions propagate upward, though interpretation requires accounting for species-specific sensitivities and confounding factors like predation.[151] Contaminant burdens in seabird tissues offer direct proxies for pollution levels, as persistent toxins like mercury and potentially toxic elements (PTEs) biomagnify through food chains, with feather and egg analyses revealing spatial gradients tied to industrial emissions and ocean circulation.[152][153] For example, North Atlantic seabirds exhibit mercury concentrations varying by latitude and foraging guild, mirroring atmospheric deposition patterns and upwelling influences.[152] Plastic ingestion, pervasive across 186 species, poses ingestion risks modeled at 99% probability for some taxa by 2050, serving as a sentinel for microplastic dispersion in surface waters.[154] Such metrics, while powerful, demand validation against direct environmental sampling to distinguish bioaccumulation from metabolic processing.[155]Nutrient Cycling and Trophic Cascades
Seabirds facilitate nutrient cycling by vectoring marine-derived nitrogen, phosphorus, and other elements to terrestrial and coastal habitats through guano, regurgitated food, eggshells, and carcasses during breeding seasons. This cross-ecosystem subsidy is substantial; modeling estimates indicate that extant seabirds transport approximately 150 million kilograms of phosphorus annually from oceans to landmasses worldwide, comparable to inputs from anadromous fish.[156] Seabird colonies act as hotspots for this deposition, with guano inputs elevating soil nitrogen and phosphorus levels by orders of magnitude in affected sites, such as desert islands where phosphorus concentrations can exceed 1,000 mg/kg compared to background levels below 100 mg/kg.[157] [158] These nutrients enhance soil fertility, microbial activity, and plant productivity; for example, in montane forests of the Pacific, endangered seabird colonies increase foliar nitrogen by 20-50% in understory vegetation, supporting denser vegetation cover and higher arthropod biomass.[159] In coastal and island ecosystems, guano solubilizes into runoff, fertilizing adjacent marine waters and boosting phytoplankton productivity by up to 30% in localized patches, as observed in sub-Antarctic studies.[160] Seabird biomass and species diversity amplify these effects, with higher-diversity colonies provisioning 2-3 times more nutrients to coral reefs and tropical islands than low-diversity ones, thereby sustaining reef-associated food webs.[161] This nutrient enrichment initiates bottom-up trophic cascades, where increased primary production cascades through herbivores and detritivores to higher trophic levels. On islands colonized by seabirds, guano-driven plant growth supports elevated invertebrate populations, which in turn fuel insectivorous vertebrates; stable isotope analyses in Aleutian seabird colonies reveal that up to 25% of terrestrial insectivore diets derive from marine subsidies, propagating productivity gains across trophic levels.[160] In marine contexts, guano plumes enhance zooplankton via algal blooms, indirectly benefiting planktivorous fish and filter feeders like manta rays, with documented increases in coral-associated biodiversity near colonies.[162] Conversely, seabirds exert top-down control as mid-to-upper trophic predators, preying on forage fish such as herring and anchovies, which can alleviate grazing pressure on zooplankton and indirectly boost primary production through reduced trophic suppression. In the Baltic Sea, reductions in cod (a shared predator) amplify sprat abundances, intensifying competition with seabirds and cascading to diminished zooplankton stocks, demonstrating how predator-prey dynamics involving seabirds propagate downward.[163] Empirical quantification remains challenging due to confounding factors like ocean currents and fisheries, but meta-analyses confirm that seabird foraging depresses prey fish densities by 10-20% locally, with knock-on effects on lower food web strata.[164] These dual mechanisms—subsidies and predation—underscore seabirds' role in stabilizing ecosystem fluxes, though anthropogenic declines in populations have attenuated these processes, reducing global nutrient transfers by an estimated 90% since pre-industrial times due to historical harvesting and habitat loss.[156]Predation and Competition Dynamics
Seabird populations experience significant predation pressure, particularly during breeding seasons when adults, eggs, and chicks are concentrated in colonies. Avian predators such as skuas (Stercorarius spp.), gulls (Larus spp.), and jaegers frequently target unattended eggs and chicks, with kleptoparasitism—stealing food from foraging adults—also common among species like the great skua (Stercorarius skua).[165] Mammalian predators, often introduced to islands, exacerbate risks; rats (Rattus spp.), cats (Felis catus), and foxes (Vulpes spp.) consume eggs and chicks, contributing to 42% of insular bird extinctions globally.[166] Empirical studies demonstrate density-dependent effects, where increased predator abundance correlates with reduced prey fecundity, as observed in systems involving yellow-legged gulls (Larus michahellis) preying on Audouin's gulls (Ichthyaetus audouinii), stabilizing populations through elevated predation rates on denser colonies.[89] In predator-prey dynamics, seabird responses include behavioral adaptations like synchronous breeding to dilute individual risk and colonial nesting for enhanced vigilance. River otters (Lontra canadensis) have been documented preying on nesting seabirds along North American coasts, with predation events peaking during chick-rearing phases.[167] Introduced avian predators, such as barn owls (Tyto alba), further intensify pressure on burrow-nesting species, altering local dynamics through direct consumption and facilitation of other predators.[168] These interactions often exhibit spatial and temporal variability, with predator activity declining during peak breeding daylight hours in some systems, potentially aligning with prey anti-predator strategies.[169] Competition among seabirds manifests primarily intraspecifically for nest sites in high-density colonies and interspecifically for marine prey resources. Aggressive territorial behaviors and eviction attempts during site selection can lead to chick mortality, with density-dependent competition influencing foraging efficiency and reproductive output.[170] Sympatric species partition foraging niches—differing in dive depths, prey sizes, or temporal patterns—to mitigate overlap, a strategy that intensifies under food scarcity, as evidenced in studies of boobies (Sula spp.) segregating by prey type and location.[171] [172] Kleptoparasitic competition, where dominant species like frigatebirds (Fregata spp.) or skuas harass others to relinquish catches, imposes energetic costs that reduce host breeding success by up to 20-30% in affected populations.[165] These predation and competition dynamics regulate seabird populations via top-down control, with empirical models showing stochastic predation driving community assembly and prey size structuring non-trophic interactions. Inter-colony competition for shared foraging grounds promotes spatial segregation, enhancing overall resilience but amplifying vulnerability when resources contract.[173] Intraspecific competition at larger scales can limit range expansions, as denser breeding aggregations face amplified risks from both endemic and invasive predators.[174]Human Interactions
Historical Harvesting and Economic Uses
Seabirds have been harvested by humans for subsistence and commercial purposes since prehistoric times, primarily for eggs, meat, feathers, and excrement used as fertilizer. In Iceland, seabird hunting and egg collection, documented in Norse sagas, formed a key subsistence resource from early settlement around 874 CE, targeting species such as puffins and guillemots with practices including cliff scaling and net traps.[175] Similarly, Indigenous groups like the Huna Tlingit in Alaska gathered gull eggs seasonally, a tradition persisting into modern regulated harvests, with over 980 eggs distributed to tribal members since 2015 under federal agreements.[176] In coastal New England, 19th-century fishermen netted nesting seabirds, salting and barreling them for market shipment, reflecting opportunistic exploitation tied to fishing economies.[177] Feathers from seabirds fueled a lucrative millinery trade in the late 19th and early 20th centuries, driving mass killings for hat decorations. Between 1897 and 1914, approximately 3.5 million seabirds, including albatrosses and petrels, were harvested in the Pacific Ocean to supply the industry, with plumes often valued higher than gold by weight.[178] This global trade targeted breeding colonies, where hunters plucked or skinned birds, leaving populations vulnerable; snowy egrets and other coastal species suffered severe declines, though seabird-specific data highlights unsustainable pressure on remote island breeders.[179] The most significant economic use involved seabird guano mining, which revolutionized 19th-century agriculture as a nitrogen-rich fertilizer. Peru's Chincha Islands, hosting massive colonies of guano-producing birds like Peruvian boobies and cormorants, yielded over 12 million tons exported from 1840 onward, generating substantial revenue and sparking international conflicts, including the U.S. Guano Islands Act of 1856 that claimed over 90 Pacific atolls.[180] By 1880, major deposits were depleted due to intensive extraction and habitat disruption, shifting reliance to synthetic alternatives, though guano's role in boosting crop yields—up to 30% in some European soils—underscored its causal impact on pre-chemical farming productivity.[181] Harvesting often employed forced labor, contributing to worker fatalities from toxic dust and collapses, while indirect effects like reduced fish availability from overfishing compounded bird declines.[182]Fisheries By-Catch and Resource Competition
Fisheries by-catch poses a significant mortality source for seabirds, primarily through entanglement in longline gear, gillnets, and trawls, with global estimates indicating 160,000 to 320,000 birds killed annually in longline fisheries alone.[183] Additional data from 2024 reveal at least 44,000 seabirds dying yearly in trawl fisheries worldwide, while gillnet by-catch may account for up to 400,000 individuals.[184] Procellariiform species, including albatrosses and petrels, suffer disproportionately, comprising over 60% of documented interactions, with hotspots in the Southern Ocean, Pacific tuna fisheries, and demersal operations off South America and Africa.[185] These incidental captures contribute to population declines in at least 20 threatened seabird taxa, exacerbating vulnerabilities in species already facing low reproductive rates.[186] Resource competition arises from spatial and dietary overlap between seabirds and commercial fisheries targeting shared prey like forage fish, small pelagics, and squid, with seabirds collectively removing a prey biomass equivalent to global commercial landings.[187] Intensified fishing pressure depletes local stocks, forcing seabirds to forage farther or switch to lower-quality prey, correlating with reduced breeding success and chick condition in colonies dependent on sardines, anchovies, and capelin.[188] Empirical studies document heightened competition in regions such as the Southern Ocean and Asian shelves, where fishery removals exceed seabird consumption, leading to measurable trophic impacts without evidence of compensatory mechanisms fully offsetting losses.[189] While discards can subsidize some scavenging species, overall fishery expansion has net negative effects on seabird demographics, as prey depletion outweighs supplemental feeding benefits.[186] Mitigation strategies for by-catch, including bird-scaring lines (tori lines), weighted branch lines, and night setting, have proven effective in reducing interactions by 70-90% when implemented in combination, as demonstrated in pelagic longline trials.[190] For instance, line weighting alone decreased by-catch by 37-76% in sablefish and cod fisheries, with further gains from integrated measures like underwater bait setters.[191] Adoption varies regionally, with mandatory regulations under frameworks like the U.S. National Plan of Action yielding declines from 6,353 seabirds in 2005 to 3,712 in 2010 in Alaska longline operations, though incomplete compliance and data gaps persist in developing-world fleets.[192] Addressing competition requires ecosystem-based fishery management to maintain forage fish quotas above thresholds supporting seabird needs, though quantifying precise allocation remains challenging due to variable seabird consumption rates.[193]Cultural and Traditional Practices
Indigenous coastal peoples in the Arctic and subarctic regions, such as the Inuit, traditionally hunted seabirds year-round using bird darts, throwing boards, snares, bows, arrows, bolas, and nets for food and materials.[194] The Unangan people of the Pribilof Islands harvested seabirds for sustenance, tools, and clothing, notably crafting renowned birdskin parkas from seabird skins.[195] In southeastern Alaska, Huna Tlingit communities annually collected glaucous-winged gull eggs from rookeries in Glacier Bay, a practice integrated into family activities and emphasizing selective harvesting to sustain populations, with only a portion of eggs taken per nest.[196][197] Chugach Alaska Native groups similarly gathered eggs from seabird islets, limiting collection to a few per nest to preserve breeding colonies.[198] In the Pacific, Rakiura Māori have conducted muttonbirding—harvesting sooty shearwater (tītī) chicks—for food, trade, and feathers since pre-European times, with the practice holding profound cultural, identity, and economic value tied to ancestral rights over specific islands. Oceanic cultures employ sustainable seabird and egg harvesting methods, often documented in oral traditions, alongside using seabirds in mythology, art, and navigation aids, such as observing white terns to locate islands during voyages.[199][200] Coastal Sámi in northern Norway maintain historical seabird utilization practices, reflecting adaptation to marine environments.[201] Seabirds feature in folklore and symbolism across cultures; albatrosses historically signified fortune and mystery for seafarers in ancient maritime tales, predating negative literary associations.[202] In Christian iconography, the pelican symbolizes piety and self-sacrifice, derived from medieval beliefs in its habit of feeding young with its blood, influencing art and heraldry from at least the 12th century.Threats and Population Trends
Empirical Drivers of Declines
Empirical studies indicate that seabird populations have experienced substantial declines globally, with an estimated 70% reduction in abundance since the 1950s, driven primarily by anthropogenic factors such as fisheries interactions, invasive predators, and altered marine food webs.[6] A comprehensive assessment of threats affecting over 170 million individual seabirds (more than 20% of the global population) highlights bycatch, invasive alien species, and habitat degradation as leading causes, with 89% of climate-impacted species also facing these overlapping pressures.[203] Bycatch in commercial fisheries, particularly pelagic longline operations targeting tuna and related species, represents a major direct mortality driver, killing hundreds of thousands of seabirds annually and contributing to population crashes in species like albatrosses and petrels.[186] A 2024 meta-analysis of standardized interaction rates across fisheries confirmed bycatch as a prominent factor in seabird declines, with observed rates varying by gear type and mitigation use, but persistent high mortality in unmitigated fleets.[204] For instance, in the Hawaii longline tuna fishery, empirical data from observer programs showed significant seabird captures prior to mandatory mitigation, correlating with regional population decreases in procellariiforms.[205] Invasive non-native predators, including rats, cats, and mongoose introduced to breeding islands, exert severe predatory pressure on ground-nesting seabirds, leading to near-total reproductive failure and colony abandonment in affected sites.[206] A global review of 115 rat-seabird interactions across 61 islands documented impacts on 75 species from 10 families, with burrowing petrels and shearwaters showing the most acute declines due to egg and chick predation.[206] Eradication efforts provide causal evidence of recovery; for example, post-removal monitoring on islands revealed rapid increases in seabird breeding success and population growth, underscoring invasives as a reversible driver distinct from broader oceanic changes.[207] Reduced prey availability from overfishing and climate-induced shifts in marine ecosystems further exacerbates declines by increasing foraging effort and lowering breeding success.[188] Studies in the North Atlantic and Alaskan waters link overexploitation of forage fish to diminished puffin and murre productivity, with empirical correlations between fishery removals and seabird chick starvation rates.[208] Climate change compounds this through ocean warming, which disrupts plankton dynamics and fish distributions, as evidenced by multi-decadal data showing productivity drops in surface-feeding seabirds across northern hemisphere systems.[209] In the Bering Sea, negative phases of climatic indices aligned with accelerated declines in ice-obligate species like least auklets, tied to sea ice loss and prey mismatches.[210] Oil pollution and plastic ingestion, while less quantified globally, demonstrate direct lethal and sublethal effects; for example, major spills have caused mass mortality events, with oiled birds exhibiting reduced insulation and foraging capacity, as observed in post-Exxon Valdez monitoring of auklets and other nearshore species.[211] These drivers interact synergistically, with fisheries depleting food resources while bycatch removes adults, amplifying vulnerability to environmental variability in long-lived, low-fecundity species.[6]Natural vs. Anthropogenic Factors
Seabird populations experience fluctuations from both natural and anthropogenic factors, though empirical assessments indicate that human-induced threats have driven the majority of long-term declines observed since the late 20th century. Natural factors primarily involve short-term variability, such as episodic prey shortages linked to oceanographic oscillations like El Niño-Southern Oscillation (ENSO) events, which reduced breeding success in species like Brandt's cormorants (Phalacrocorax penicillatus) along the California Current by disrupting forage fish availability in the early 1980s.[212] Disease outbreaks and intrinsic density-dependent regulation also contribute to natural mortality, but these rarely cause sustained population crashes without amplification by external pressures.[213] Native predation, while present in some ecosystems, is typically balanced by evolutionary adaptations in seabirds that nest on predator-free islands or cliffs.[214] In contrast, anthropogenic factors exert persistent, compounding effects that override natural resilience. Introduced invasive predators, such as rats (Rattus spp.) and cats (Felis catus), introduced by human activity, have decimated breeding colonies by preying on eggs and chicks; for instance, eradication efforts on islands have led to rapid population recoveries in affected species, demonstrating direct causality.[215] Fisheries bycatch remains a leading marine threat, with longline and gillnet fisheries entangling and drowning millions of seabirds annually, particularly albatrosses and petrels, as evidenced by global tracking data showing overlap between foraging ranges and fishing grounds.[203] Overfishing depletes prey stocks, exacerbating food competition, while pollution from plastics and oil causes chronic mortality; ingested plastics impair reproduction, and oil spills, like the 1989 Exxon Valdez incident, killed tens of thousands of birds through hypothermia and toxicity.[216][203] Comparative analyses reveal that while natural climate variability induces cyclical booms and busts—such as puffin (Fratercula arctica) breeding failures during local prey shortages—anthropogenic drivers like bycatch and invasives correlate with irreversible declines, affecting over 30% of seabird species classified as threatened.[208][217] Interventions targeting human factors, including predator removal and bycatch mitigation via gear modifications, have stabilized or increased populations in targeted areas, underscoring their outsized role over natural processes.[218] For example, a global review estimates that addressing invasives, bycatch, and overfishing could benefit 380 million individual seabirds, far exceeding gains from managing natural variability alone.[203] This distinction highlights the need for causal attribution based on demographic modeling and intervention outcomes rather than correlative associations often amplified in environmental narratives.[219]Global and Regional Trend Data
Global seabird populations have experienced substantial declines, with approximately 50% of the 369 recognized species showing decreasing trends over the past 50 years and an estimated overall population reduction of 70%.[211] Analysis of monitored populations, representing about 19% of the global total and drawn from 9,920 records across 3,213 breeding sites, indicates a 69.7% decline from 1950 to 2010, with the steepest drops in families such as terns (85.8%) and procellariids (79.6%).[147] As of the latest IUCN assessments, 30% of seabird species are classified as threatened (Critically Endangered, Endangered, or Vulnerable), and 11% as Near Threatened, reflecting ongoing pressures despite some stable or locally increasing populations.[211] In Europe, encompassing 80 seabird species, 34% exhibit decreasing population trends, with 32% categorized as threatened or Near Threatened.[211] The 2023 Seabirds Count census for Britain and Ireland revealed that 11 of 21 monitored species with reliable trend data had declined by more than 10% since the prior census around 2000, including sharp drops in kittiwakes (up to 43% in some areas) and Arctic terns.[220] Within the European Union, 38% of 66 assessed seabird species show declines, with notable uplistings such as the Greater Scaup to Endangered due to rapid reductions.[211] Regional variations highlight differential impacts across ocean basins. In the North Atlantic and Baltic Sea, breeding abundances of several species, such as common murres and black-legged kittiwakes, have declined more severely than in adjacent North Sea populations, with Finnish coastal trends showing steeper drops linked to local environmental indicators.[221] Southern Hemisphere examples include significant reductions in sooty terns in French Polynesia and guanay cormorants off Peru, contributing to broader pelagic family declines.[147] A proposed productivity-based indicator for northern European seabirds estimates current breeding success could sustain annual declines of 3-4%.[222]| Region/Ocean Basin | Key Trend Observations | Example Species Declines |
|---|---|---|
| Europe (Pan-European) | 34% of species decreasing; 32% threatened/NT | Balearic Shearwater (Critically Endangered uplisting); Northern Fulmar (Vulnerable)[211] |
| North Atlantic/Baltic | Steeper declines vs. North Sea; productivity-driven | Common murre, black-legged kittiwake[221] |
| Southern Hemisphere (e.g., Pacific/Peru) | Major pelagic losses 1950-2010 | Sooty tern (French Polynesia), guanay cormorant[147] |
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