Runde is an island in the municipality of Herøy in Møre og Romsdal county, Norway . The island has a population of 113 people (as of 2015), and it is connected by the Runde Bridge to the island of Remøya to the south.
133-449: Runde is famous for its enormous number of birds. It is said that there are about 500,000 to 700,000 seabirds inhabiting the island, most of them are to be found in the cliffs. The Runde Environmental Centre ( Norwegian : Runde Miljøsenter ) is a research station at the bird island. Runde lies on the west coast of Norway, close to towns and villages like Fosnavåg , Ulsteinvik , Volda , Ørsta , and Ålesund . This region of Norway
266-409: A Gaussian might describe the frequency with which a species ate prey of a certain size, giving a more detailed niche description than simply specifying some median or average prey size. For such a bell-shaped distribution, the position , width and form of the niche correspond to the mean , standard deviation and the actual distribution itself. One advantage in using statistics is illustrated in
399-447: 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
532-447: A storage effect . Species can differentiate their niche via a competition-predation trade-off if one species is a better competitor when predators are absent, and the other is better when predators are present. Defenses against predators, such as toxic compounds or hard shells, are often metabolically costly. As a result, species that produce such defenses are often poor competitors when predators are absent. Species can coexist through
665-401: A wreck . Seabirds have had a long association with both fisheries and sailors , and both have drawn benefits and disadvantages from the relationship. Fishermen have traditionally used seabirds as indicators of both fish shoals , underwater banks that might indicate fish stocks, and of potential landfall. In fact, the known association of seabirds with land was instrumental in allowing
798-727: A certain environment (have overlapping requirement niches) but fundamentally differ the ways that they use (or "impact") that environment. These requirements have repeatedly been violated by nonnative (i.e. introduced and invasive ) species, which often coexist with new species in their nonnative ranges, but do not appear to be constricted these requirements. In other words, contemporary niche theory predicts that species will be unable to invade new environments outside of their requirement (i.e. realized) niche, yet many examples of this are well-documented. Additionally, contemporary niche theory predicts that species will be unable to establish in environments where other species already consume resources in
931-618: A clade, the Aequornithes either became seabirds in a single transition in the Cretaceous or some lineages such as pelicans and frigatebirds adapted to sea living independently from freshwater-dwelling ancestors. In the Paleogene both pterosaurs and marine reptiles became extinct, allowing seabirds to expand ecologically. These post-extinction seas were dominated by early Procellariidae , giant penguins and two extinct families ,
1064-427: A climatic perspective, to explain distribution and abundance. Current predictions on species responses to climate change strongly rely on projecting altered environmental conditions on species distributions. However, it is increasingly acknowledged that climate change also influences species interactions and an Eltonian perspective may be advantageous in explaining these processes. This perspective of niche allows for
1197-543: 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. 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
1330-482: A competition-predation trade-off if predators are more abundant when the less defended species is common, and less abundant if the well-defended species is common. This effect has been criticized as being weak, because theoretical models suggest that only two species within a community can coexist because of this mechanism. Two ecological paradigms deal with the problem. The first paradigm predominates in what may be called "classical" ecology. It assumes that niche space
1463-486: A fashion similar to grebes and loons (using its feet to move underwater) but had a beak filled with sharp teeth. Flying Cretaceous seabirds do not exceed wingspans of two meters; any sizes were taken by piscivorous pterosaurs . While Hesperornis is not thought to have left descendants, the earliest modern seabirds also occurred in the Cretaceous, with a species called Tytthostonyx glauconiticus , which has features suggestive of Procellariiformes and Fregatidae. As
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#17327811383151596-472: A food source for predators and a consumer of prey). "The type and number of variables comprising the dimensions of an environmental niche vary from one species to another [and] the relative importance of particular environmental variables for a species may vary according to the geographic and biotic contexts". A Grinnellian niche is determined by the habitat in which a species lives and its accompanying behavioral adaptations . An Eltonian niche emphasizes that
1729-412: A fundamental niche of the entire slope of a hillside, but its realized niche is only the top portion of the slope because species Y, which is a better competitor but cannot survive on the top portion of the slope, has excluded it from the lower portion of the slope. With this scenario, competition will continue indefinitely in the middle of the slope between these two species. Because of this, detection of
1862-480: A given species), 'niche partitioning' (resource differentiation by coexisting species), and 'niche overlap' (overlap of resource use by different species). Statistics were introduced into the Hutchinson niche by Robert MacArthur and Richard Levins using the 'resource-utilization' niche employing histograms to describe the 'frequency of occurrence' as a function of a Hutchinson coordinate. So, for instance,
1995-971: A large number of non-governmental organizations (including BirdLife International , the American Bird Conservancy and the Royal Society for the Protection of Birds ). 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). Many seabirds are little studied and poorly known because they live far out at sea and breed in isolated colonies. Some seabirds, particularly
2128-570: A lineage— Eurypygimorphae —that is a sister group to the Aequornithes. Seabirds, by virtue of living in a geologically depositional environment (that is, in the sea where sediments are readily laid down), are well represented in the fossil record. They are first known to occur in the Cretaceous period, the earliest being the Hesperornithiformes , like Hesperornis regalis , a flightless loon-like seabird that could dive in
2261-587: A million birds have been recorded, both in the tropics (such as Kiritimati in the Pacific ) and in the polar latitudes (as in Antarctica ). Seabird colonies occur exclusively for the purpose of breeding; non-breeding birds will only collect together outside the breeding season in areas where prey species are densely aggregated. Seabird colonies are highly variable. Individual nesting sites can be widely spaced, as in an albatross colony, or densely packed as with
2394-541: A new legislation, making all wrecks older than 100 years automatically protected in Norway. The Runde Environmental Centre is a research station on Runde which has 4 main activities: a marine station, a house and hub for small spin-off businesses, an information centre, and a conference/overnight centre with 34 hotel-standard rooms in 8 apartment suites. In 2008, the Runde Tourist office at Runde Environmental Centre
2527-465: A niche specific to each species. Species can however share a 'mode of life' or 'autecological strategy' which are broader definitions of ecospace. For example, Australian grasslands species, though different from those of the Great Plains grasslands, exhibit similar modes of life. Once a niche is left vacant, other organisms can fill that position. For example, the niche that was left vacant by
2660-693: 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. 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
2793-414: A place for returning mates to reunite, and reduces the costs of prospecting for a new site. Young adults breeding for the first time usually return to their natal colony, and often nest close to where they hatched. This tendency, known as philopatry , is so strong that a study of Laysan albatrosses found that the average distance between hatching site and the site where a bird established its own territory
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#17327811383152926-438: A reduced capacity for powered flight and are dependent on a type of gliding called dynamic soaring (where the wind deflected by waves provides lift) as well as slope soaring. Seabirds also almost always have webbed feet , to aid movement on the surface as well as assisting diving in some species. The Procellariiformes are unusual among birds in having a strong sense of smell , which is used to find widely distributed food in
3059-461: A smaller layer of air (compared to other diving birds) but otherwise soak up water. This allows them to swim without fighting the buoyancy that retaining air in the feathers causes, yet retain enough air to prevent the bird losing excessive heat through contact with water. The plumage of most seabirds is less colourful than that of land birds, restricted in the main to variations of black, white or grey. A few species sport colourful plumes (such as
3192-575: 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. 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. Changes to
3325-460: A species not only grows in and responds to an environment, it may also change the environment and its behavior as it grows. The Hutchinsonian niche uses mathematics and statistics to try to explain how species coexist within a given community. The concept of ecological niche is central to ecological biogeography , which focuses on spatial patterns of ecological communities. "Species distributions and their dynamics over time result from properties of
3458-487: A species' density declines, then the food it most depends on will become more abundant (since there are so few individuals to consume it). As a result, the remaining individuals will experience less competition for food. Although "resource" generally refers to food, species can partition other non-consumable objects, such as parts of the habitat. For example, warblers are thought to coexist because they nest in different parts of trees. Species can also partition habitat in
3591-403: A vast ocean, and help distinguish familiar nest odours from unfamiliar ones. Salt glands are used by seabirds to deal with the salt they ingest by drinking and feeding (particularly on crustaceans ), and to help them osmoregulate . The excretions from these glands (which are positioned in the head of the birds, emerging from the nasal cavity ) are almost pure sodium chloride . With
3724-400: A very variable prey source); this may be a reason why it arises more frequently in seabirds. There are other possible advantages: colonies may act as information centres, where seabirds returning to the sea to forage can find out where prey is by studying returning individuals of the same species. There are disadvantages to colonial life, particularly the spread of disease. Colonies also attract
3857-451: A way that gives them access to different types of resources. As stated in the introduction, anole lizards appear to coexist because each uses different parts of the forests as perch locations. This likely gives them access to different species of insects. Research has determined that plants can recognize each other's root systems and differentiate between a clone, a plant grown from the same mother plants seeds, and other species. Based on
3990-458: A year, unless they lose the first (with a few exceptions, like the Cassin's auklet ), and many species (like the tubenoses and sulids ) will only lay one egg a year. Care of young is protracted, extending for as long as six months, among the longest for birds. For example, once common guillemot chicks fledge , they remain with the male parent for several months at sea. The frigatebirds have
4123-511: Is absent or low, and therefore detection of niche differentiation is difficult or impossible. Below is a list of ways that species can partition their niche. This list is not exhaustive, but illustrates several classic examples. Resource partitioning is the phenomenon where two or more species divides out resources like food, space, resting sites etc. to coexist. For example, some lizard species appear to coexist because they consume insects of differing sizes. Alternatively, species can coexist on
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4256-548: Is constrained by different natural enemies, they will be able to coexist. Early work focused on specialist predators; however, more recent studies have shown that predators do not need to be pure specialists, they simply need to affect each prey species differently. The Janzen–Connell hypothesis represents a form of predator partitioning. Conditional differentiation (sometimes called temporal niche partitioning ) occurs when species differ in their competitive abilities based on varying environmental conditions. For example, in
4389-407: 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. Gannets , boobies , tropicbirds , some terns, and brown pelicans all engage in plunge diving, taking fast-moving prey by diving into the water from the flight. Plunge diving allows birds to use the energy from the momentum of
4522-730: Is famous for its long fjords and steep, snow-clad mountains. The island has been designated an Important Bird Area (IBA) by BirdLife International . The birdlife on Runde is protected by the Goksøyr Mires Nature Reserve , part of the Runde Ramsar Site . Throughout the years, several ships have sunk near Runde on the Norwegian coast. Some of them are said to be ships from the Netherlands and Spain , loaded with gold and silver . One ship from
4655-481: Is greater than inter-specific (between species) competition. Since niche differentiation concentrates competition within-species, due to a decrease in between-species competition, the Lotka-Volterra model predicts that niche differentiation of any degree will result in coexistence. In reality, this still leaves the question of how much differentiation is needed for coexistence. A vague answer to this question
4788-642: Is inconclusive. Some plunge divers (as well as some surface feeders) are dependent on dolphins and tuna to push shoaling fish up towards the surface. This catch-all category refers to other seabird strategies that involve the next trophic level up. Kleptoparasites are seabirds that make a part of their living stealing food of other seabirds. Most famously, frigatebirds and skuas engage in this behaviour, although gulls, terns and other species will steal food opportunistically. The nocturnal nesting behaviour of some seabirds has been interpreted as arising due to pressure from this aerial piracy. Kleptoparasitism
4921-489: Is largely saturated with individuals and species, leading to strong competition. Niches are restricted because "neighbouring" species, i.e., species with similar ecological characteristics such as similar habitats or food preferences, prevent expansion into other niches or even narrow niches down. This continual struggle for existence is an important assumption of natural selection introduced by Darwin as an explanation for evolution. The other paradigm assumes that niche space
5054-455: Is merely the product of the competitive exclusion principle. Also, because no species is out-competing any other species in the final community, the presence of niche differentiation will be difficult or impossible to detect. Finally, niche differentiation can arise as an evolutionary effect of competition. In this case, two competing species will evolve different patterns of resource use so as to avoid competition. Here too, current competition
5187-405: Is necessary for ecologists to be able to detect, measure, and quantify the niches of different coexisting and competing species. This is often done through a combination of detailed ecological studies, controlled experiments (to determine the strength of competition), and mathematical models . To understand the mechanisms of niche differentiation and competition, much data must be gathered on how
5320-545: Is not thought to play a significant part of the diet of any species, and is instead a supplement to food obtained by hunting. A study of great frigatebirds stealing from masked boobies estimated that the frigatebirds could at most obtain 40% of the food they needed, and on average obtained only 5%. Many species of gull will feed on seabird and sea mammal carrion when the opportunity arises, as will giant petrels . Some species of albatross also engage in scavenging: an analysis of regurgitated squid beaks has shown that many of
5453-402: 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. 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
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5586-646: 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. Ecological niche In ecology , a niche is the match of a species to a specific environmental condition. It describes how an organism or population responds to the distribution of resources and competitors (for example, by growing when resources are abundant, and when predators , parasites and pathogens are scarce) and how it in turn alters those same factors (for example, limiting access to resources by other organisms, acting as
5719-489: Is some evidence of this, the effects of seabirds are considered smaller than that of marine mammals and predatory fish (like tuna ). 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. This can have other impacts; for example,
5852-419: Is that the more similar two species are, the more finely balanced the suitability of their environment must be in order to allow coexistence. There are limits to the amount of niche differentiation required for coexistence, and this can vary with the type of resource, the nature of the environment, and the amount of variation both within and between the species. To answer questions about niche differentiation, it
5985-783: Is that they feed in saltwater ; but, as seems to be true with any statement in biology, some do not." However, by convention all of the Sphenisciformes (penguins) and Procellariiformes ( albatrosses and petrels ), all of the Suliformes ( gannets and cormorants ) except the darters , and some of the Charadriiformes (the 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
6118-589: Is the skimmer , which has a unique fishing method: flying along the surface with the lower mandible in the water—this shuts automatically when the bill touches something in the water. The skimmer's bill reflects its unusual lifestyle, with the lower mandible uniquely being longer than the upper one. Surface feeders that swim often have unique bills as well, adapted for their specific prey. Prions have special bills with filters called lamellae to filter out plankton from mouthfuls of water, and many albatrosses and petrels have hooked bills to snatch fast-moving prey. On
6251-448: Is the deepest diver of the shearwaters, having been recorded diving below 70 metres (230 ft). Some albatross species are also capable of limited diving, with light-mantled sooty albatrosses holding the record at 12 metres (40 ft). 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
6384-425: Is the niche that is filled by birds of prey which eat small animals such as shrews and mice. In an oak wood this niche is filled by tawny owls , while in the open grassland it is occupied by kestrels . The existence of this carnivore niche is dependent on the further fact that mice form a definite herbivore niche in many different associations, although the actual species of mice may be quite different. Conceptually,
6517-432: Is to a large degree vacant, i.e., that there are many vacant niches . It is based on many empirical studies and theoretical investigations especially of Kauffman 1993. Causes of vacant niches may be evolutionary contingencies or brief or long-lasting environmental disturbances. Both paradigms agree that species are never "universal" in the sense that they occupy all possible niches; they are always specialized, although
6650-571: 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
6783-715: The Pelagornithidae and the Plotopteridae (a group of large seabirds that looked like the penguins). Modern genera began their wide radiation in the Miocene , although the genus Puffinus (which includes today's Manx shearwater and sooty shearwater ) might date back to the Oligocene . Within the Charadriiformes, the gulls and allies ( Lari ) became seabirds in the late Eocene, and then waders in
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#17327811383156916-703: The Polynesians to locate tiny landmasses in the Pacific. Seabirds have provided food for fishermen away from home, as well as bait. Famously, tethered cormorants have been used to catch fish directly. Indirectly, fisheries have also benefited from guano from colonies of seabirds acting as fertilizer for the surrounding seas. Negative effects on fisheries are mostly restricted to raiding by birds on aquaculture , although long-lining fisheries also have to deal with bait stealing. There have been claims of prey depletion by seabirds of fishery stocks, and while there
7049-553: The Sonoran Desert , some annual plants are more successful during wet years, while others are more successful during dry years. As a result, each species will have an advantage in some years, but not others. When environmental conditions are most favorable, individuals will tend to compete most strongly with member of the same species. For example, in a dry year, dry-adapted plants will tend to be most limited by other dry-adapted plants. This can help them to coexist through
7182-677: 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. 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
7315-419: The razorbill (an Atlantic auk) requires 64% more energy to fly than a petrel of equivalent size. 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
7448-481: The shearwaters and gadfly petrels). Surface feeders in flight include some of the most acrobatic of seabirds, which either snatch morsels from the water (as do frigate-birds and some terns), or "walk", pattering and hovering on the water's surface, as some of the storm-petrels do. Many of these do not ever land in the water, and some, such as the frigatebirds, have difficulty getting airborne again should they do so. Another seabird family that does not land while feeding
7581-478: The "impact niche" is defined as the combination of effects that a given consumer has on both a). the resources that it uses, and b). the other consumers in the ecosystem. Therefore, the impact niche is equivalent to the Eltonian niche since both concepts are defined by the impact of a given species on its environment. The range of environmental conditions where a species can successfully survive and reproduce (i.e.
7714-526: The Antarctic mainland, are unlikely to find anything to eat around their breeding sites. The marbled murrelet nests inland in old growth forest , seeking huge conifers with large branches to nest on. Other species, such as the California gull , nest and feed inland on lakes, and then move to the coasts in the winter. Some cormorant, pelican , gull and tern species have individuals that never visit
7847-642: 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). 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
7980-726: The Austral summer in Antarctica. Other species also undertake trans-equatorial trips, both from the north to the south, and from south to north. The population of elegant terns , which nest off Baja California , splits after the breeding season with some birds travelling north to the Central Coast of California and some travelling as far south as Peru and Chile to feed in the Humboldt Current . The sooty shearwater undertakes an annual migration cycle that rivals that of
8113-495: The Eltonian niche introduces the idea of a species' response to and effect on the environment. Unlike other niche concepts, it emphasizes that a species not only grows in and responds to an environment based on available resources, predators, and climatic conditions, but also changes the availability and behavior of those factors as it grows. In an extreme example, beavers require certain resources in order to survive and reproduce, but also construct dams that alter water flow in
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#17327811383158246-574: The Hutchinsonian definition of a realized niche) is also encompassed under contemporary niche theory, termed the "requirement niche". The requirement niche is bounded by both the availability of resources as well as the effects of coexisting consumers (e.g. competitors and predators). Contemporary niche theory provides three requirements that must be met in order for two species (consumers) to coexist: These requirements are interesting and controversial because they require any two species to share
8379-711: The Netherlands, the Akerendam , was a newly built vessel which set sails from an island called Texel in Netherlands on 19 January 1725. Akerendam was a part of the Dutch merchant fleet, headed for Batavia ( Indonesia ) loaded with gold and silver coins to be used for the trade of spices in the Far East . The ship got caught in a storm in the North Sea , and the Akerendam disappeared into the north. The ship went down on
8512-522: The Norwegian Coin Museum in Oslo . The finders were allowed to keep two thirds of the treasure, which was later sold to collectors worldwide. The 1724 Utrecht Gold Ducats can command $ 750 to $ 1,000 each and are all in uncirculated quality, the highest graded ones are NGC / PCGS MS-63. The Norwegian state received 25% and the Netherlands received 7% of the treasure. This event later initiated
8645-556: 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
8778-480: 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. For example, the northern royal albatross colony at Taiaroa Head in New Zealand attracts 40,000 visitors a year. 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
8911-467: The albatrosses and gulls, are more well known to humans. The albatross has been described as "the most legendary of birds", 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 that derives from Samuel Taylor Coleridge 's famous poem, " The Rime of the Ancient Mariner ", in which a sailor
9044-452: The attention of predators , principally other birds, and many species attend their colonies nocturnally to avoid predation. Birds from different colonies often forage in different areas to avoid competition. 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
9177-454: The bird is young. After fledging, juvenile birds often disperse further than adults, and to different areas, so are commonly sighted far from a species' normal range. Some species, such as the auks, do not have a concerted migration effort, but drift southwards as the winter approaches. Other species, such as some of the storm petrels, diving petrels and cormorants, never disperse at all, staying near their breeding colonies year round. While
9310-486: 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. In Greenland , however, uncontrolled hunting is pushing many species into steep decline. Other human factors have led to declines and even extinctions in seabird populations and species. Of these, perhaps
9443-441: The coordinate system." The niche concept was popularized by the zoologist G. Evelyn Hutchinson in 1957. Hutchinson inquired into the question of why there are so many types of organisms in any one habitat. His work inspired many others to develop models to explain how many and how similar coexisting species could be within a given community, and led to the concepts of 'niche breadth' (the variety of resources or habitats used by
9576-453: 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
9709-451: The disease have scarred digestive tracts from ingesting plastic waste . "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." The threats faced by seabirds have not gone unnoticed by scientists or the conservation movement . As early as 1903, U.S. President Theodore Roosevelt
9842-456: The dive to combat natural buoyancy (caused by air trapped in plumage), and thus uses less energy than the dedicated pursuit divers, allowing them to utilise more widely distributed food resources, for example, in impoverished tropical seas. In general, this is the most specialised method of hunting employed by seabirds; other non-specialists (such as gulls and skuas) may employ it but do so with less skill and from lower heights. In brown pelicans,
9975-413: The earliest instances known is in southern Chile, where archaeological excavations in middens has shown hunting of albatrosses, cormorants and shearwaters from 5000 BP. 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
10108-481: The ecological space occupied by a species) is subtly different from the "niche" as defined by Grinnell (an ecological role, that may or may not be actually filled by a species—see vacant niches ). A niche is a very specific segment of ecospace occupied by a single species. On the presumption that no two species are identical in all respects (called Hardin's 'axiom of inequality' ) and the competitive exclusion principle , some resource or adaptive dimension will provide
10241-801: The environment. As an example of niche partitioning, several anole lizards in the Caribbean islands share common diets—mainly insects. They avoid competition by occupying different physical locations. Although these lizards might occupy different locations, some species can be found inhabiting the same range, with up to 15 in certain areas. For example, some live on the ground while others are arboreal. Species who live in different areas compete less for food and other resources, which minimizes competition between species. However, species who live in similar areas typically compete with each other. The Lotka–Volterra equation states that two competing species can coexist when intra-specific (within species) competition
10374-463: The estimation of the competition coefficients. This postulate, however, can be misguided, as it ignores the impacts that the resources of each category have on the organism and the impacts that the organism has on the resources of each category. For instance, the resource in the overlap region can be non-limiting, in which case there is no competition for this resource despite niche overlap. An organism free of interference from other species could use
10507-404: 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
10640-587: The existence of both ecological equivalents and empty niches. An ecological equivalent to an organism is an organism from a different taxonomic group exhibiting similar adaptations in a similar habitat, an example being the different succulents found in American and African deserts, cactus and euphorbia , respectively. As another example, the anole lizards of the Greater Antilles are a rare example of convergent evolution , adaptive radiation , and
10773-482: The existence of ecological equivalents: the anole lizards evolved in similar microhabitats independently of each other and resulted in the same ecomorphs across all four islands. In 1927 Charles Sutherland Elton , a British ecologist , defined a niche as follows: "The 'niche' of an animal means its place in the biotic environment, its relations to food and enemies ." Elton classified niches according to foraging activities ("food habits"): For instance there
10906-472: The exotic or invasive species . The mathematical representation of a species' fundamental niche in ecological space, and its subsequent projection back into geographic space, is the domain of niche modelling . Contemporary niche theory (also called "classic niche theory" in some contexts) is a framework that was originally designed to reconcile different definitions of niches (see Grinnellian, Eltonian, and Hutchinsonian definitions above), and to help explain
11039-427: The extinction of the tarpan has been filled by other animals (in particular a small horse breed, the konik ). Also, when plants and animals are introduced into a new environment, they have the potential to occupy or invade the niche or niches of native organisms, often outcompeting the indigenous species. Introduction of non-indigenous species to non-native habitats by humans often results in biological pollution by
11172-676: The feathers resist abrasion. 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
11305-434: The figure, where it is clear that for the narrower distributions (top) there is no competition for prey between the extreme left and extreme right species, while for the broader distribution (bottom), niche overlap indicates competition can occur between all species. The resource-utilization approach postulates that not only can competition occur, but that it does occur, and that overlap in resource utilization directly enables
11438-475: The first time in over a hundred years. 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, and their deployment is increasingly required by many national fishing fleets. One of the Millennium Projects in
11571-478: The frequency of breeding failures due to unfavourable marine conditions, and the relative lack of predation compared to that of land-living birds. Because of the greater investment in raising the young and because foraging for food may occur far from the nest site, in all seabird species except the phalaropes, both parents participate in caring for the young, and pairs are typically at least seasonally monogamous . Many species, such as gulls, auks and penguins, retain
11704-433: The full range of conditions (biotic and abiotic) and resources in which it could survive and reproduce which is called its fundamental niche . However, as a result of pressure from, and interactions with, other organisms (i.e. inter-specific competition) species are usually forced to occupy a niche that is narrower than this, and to which they are mostly highly adapted ; this is termed the realized niche . Hutchinson used
11837-401: The idea of competition for resources as the primary mechanism driving ecology, but overemphasis upon this focus has proved to be a handicap for the niche concept. In particular, overemphasis upon a species' dependence upon resources has led to too little emphasis upon the effects of organisms on their environment, for instance, colonization and invasions. The term "adaptive zone" was coined by
11970-399: The idea that the niche of a species is determined by the habitat in which it lives and its accompanying behavioral adaptations . In other words, the niche is the sum of the habitat requirements and behaviors that allow a species to persist and produce offspring. For example, the behavior of the California thrasher is consistent with the chaparral habitat it lives in—it breeds and feeds in
12103-468: The longest period of parental care of any bird except a few raptors and the southern ground hornbill , with each chick fledging after four to six months and continued assistance after that for up to fourteen months. Due to the extended period of care, breeding occurs every two years rather than annually for some species. This life-history strategy has probably evolved both in response to the challenges of living at sea (collecting widely scattered prey items),
12236-408: The marine ecosystems caused by dredging, which alters the biodiversity of the seafloor, can also have a negative impact. 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
12369-542: The meaning of niche as a recess in a wall for a statue, which itself is probably derived from the Middle French word nicher , meaning to nest . The term was coined by the naturalist Roswell Hill Johnson but Joseph Grinnell was probably the first to use it in a research program in 1917, in his paper "The niche relationships of the California Thrasher". The Grinnellian niche concept embodies
12502-580: The middle Miocene ( Langhian ). 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. Seabirds have made numerous adaptations to living on and feeding in
12635-490: The most desirable nesting spaces. The tropical Bonin petrel nests during the winter to avoid competition with the more aggressive wedge-tailed shearwater . When the seasons overlap, the wedge-tailed shearwaters will kill young Bonin petrels in order to use their burrows. Many seabirds show remarkable site fidelity , returning to the same burrow, nest or site for many years, and they will defend that site from rivals with great vigour. This increases breeding success, provides
12768-552: 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. 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. The disturbance of breeding colonies by humans
12901-475: The narrow extent of focus, data sets characterizing Eltonian niches typically are in the form of detailed field studies of specific individual phenomena, as the dynamics of this class of niche are difficult to measure at a broad geographic scale. However, the Eltonian niche may be useful in the explanation of a species' endurance of global change. Because adjustments in biotic interactions inevitably change abiotic factors, Eltonian niches can be useful in describing
13034-426: The north side of Runde, and the whole crew of 200 was lost at sea. The inhabitants of Runde started to find parts of the ship that was washed up at its shore, including dead crew members. The wreck site was close to the shore, and during the summer of 1725 four chests and other parts of the cargo were salvaged. But then the divers gave up, and the wreck was forgotten. During the 19th century, locals often found coins at
13167-428: 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. Some seabirds have used changing wind patterns to forage further and more efficiently. In 2023, plasticosis , a new disease caused solely by plastics, was discovered in seabirds. The birds identified as having
13300-444: The other closely related species within the same broad taxonomic class, but there are exceptions. A premier example of a non-standard niche filling species is the flightless, ground-dwelling kiwi bird of New Zealand, which feeds on worms and other ground creatures, and lives its life in a mammal-like niche. Island biogeography can help explain island species and associated unfilled niches. The ecological meaning of niche comes from
13433-841: The other hand, most gulls are versatile and opportunistic feeders who will eat a wide variety of prey, both at sea and on land. Pursuit diving exerts greater pressures (both evolutionary and physiological) on seabirds, but the reward is a greater area in which to feed than is available to surface feeders. Underwater propulsion is provided by wings (as used by penguins, auks, diving petrels and some other species of petrel) or feet (as used by cormorants, grebes , loons and several types of fish-eating ducks ). Wing-propelled divers are generally faster than foot-propelled divers. The use of wings or feet for diving has limited their utility in other situations: loons and grebes walk with extreme difficulty (if at all), penguins cannot fly, and auks have sacrificed flight efficiency in favour of diving. For example,
13566-402: The other to extinction. This rule also states that two species cannot occupy the same exact niche in a habitat and coexist together, at least in a stable manner. When two species differentiate their niches, they tend to compete less strongly, and are thus more likely to coexist. Species can differentiate their niches in many ways, such as by consuming different foods, or using different areas of
13699-561: The overall response of a species to new environments. The Hutchinsonian niche is an " n-dimensional hypervolume", where the dimensions are environmental conditions and resources , that define the requirements of an individual or a species to practice its way of life, more particularly, for its population to persist. The "hypervolume" defines the multi-dimensional space of resources (e.g., light, nutrients, structure, etc.) available to (and specifically used by) organisms, and "all species other than those under consideration are regarded as part of
13832-441: The paleontologist George Gaylord Simpson to explain how a population could jump from one niche to another that suited it, jump to an 'adaptive zone', made available by virtue of some modification, or possibly a change in the food chain , that made the adaptive zone available to it without a discontinuity in its way of life because the group was 'pre-adapted' to the new ecological opportunity. Hutchinson's "niche" (a description of
13965-410: The past, several species inhabited an area, and all of these species had overlapping fundamental niches. However, through competitive exclusion, the less competitive species were eliminated, leaving only the species that were able to coexist (i.e. the most competitive species whose realized niches did not overlap). Again, this process does not include any evolutionary change of individual species, but it
14098-454: The presence of niche differentiation (through competition) will be relatively easy. Importantly, there is no evolutionary change of the individual species in this case; rather this is an ecological effect of species Y out-competing species X within the bounds of species Y's fundamental niche. Another way by which niche differentiation can arise is via the previous elimination of species without realized niches. This asserts that at some point in
14231-503: 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 , 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
14364-565: The river where the beaver lives. Thus, the beaver affects the biotic and abiotic conditions of other species that live in and near the watershed. In a more subtle case, competitors that consume resources at different rates can lead to cycles in resource density that differ between species. Not only do species grow differently with respect to resource density, but their own population growth can affect resource density over time . Eltonian niches focus on biotic interactions and consumer–resource dynamics (biotic variables) on local scales. Because of
14497-658: The root meristem in that direction, if the interaction is kin. Simonsen discusses how plants accomplish root communication with the addition of beneficial rhizobia and fungal networks and the potential for different genotypes of the kin plants, such as the legume M. Lupulina, and specific strains of nitrogen fixing bacteria and rhizomes can alter relationships between kin and non-kin competition. This means there could be specific subsets of genotypes in kin plants that selects well with specific strains that could outcompete other kin. What might seem like an instance in kin competition could just be different genotypes of organisms at play in
14630-403: The root secretions, also called exudates, plants can make this determination. The communication between plants starts with the secretions from plant roots into the rhizosphere. If another plant that is kin is entering this area the plant will take up exudates. The exudate, being several different compounds, will enter the plants root cell and attach to a receptor for that chemical halting growth of
14763-465: The same mate for several seasons, and many petrel species mate for life. Albatrosses and procellariids , which mate for life, take many years to form a pair bond before they breed, and the albatrosses have an elaborate breeding dance that is part of pair-bond formation. Ninety-five percent of seabirds are colonial, and seabird colonies are among the largest bird colonies in the world, providing one of Earth's great wildlife spectacles. Colonies of over
14896-484: The same resources if each species is limited by different resources, or differently able to capture resources. Different types of phytoplankton can coexist when different species are differently limited by nitrogen, phosphorus, silicon, and light. In the Galapagos Islands , finches with small beaks are more able to consume small seeds, and finches with large beaks are more able to consume large seeds. If
15029-472: The same ways as the incoming species, however examples of this are also numerous. In ecology , niche differentiation (also known as niche segregation , niche separation and niche partitioning ) refers to the process by which competing species use the environment differently in a way that helps them to coexist. The competitive exclusion principle states that if two species with identical niches (ecological roles) compete , then one will inevitably drive
15162-653: The sea at all, spending their lives on lakes, rivers, swamps and, in the case of some of the gulls, cities and agricultural land. In these cases, it is thought that these terrestrial or freshwater birds evolved from marine ancestors. Some seabirds, principally those that nest in tundra , as skuas and phalaropes do, will migrate over land as well. The more marine species, such as petrels, auks and gannets , are more restricted in their habits, but are occasionally seen inland as vagrants. This most commonly happens to young inexperienced birds, but can happen in great numbers to exhausted adults after large storms , an event known as
15295-507: The sea's edge (coast), but are also not treated as seabirds. Sea eagles and other fish-eating birds of prey are also typically excluded, however tied to marine environments they may be. German ornithologist Gerald Mayr defined the "core waterbird" clade Aequornithes in 2010. This lineage gives rise to the Gaviiformes , Sphenisciformes , Procellariiformes, Ciconiiformes , Suliformes and Pelecaniformes . The tropicbirds are part of
15428-407: The sea. Wing morphology has been shaped by the niche an individual species or family has evolved , so that looking at a wing's shape and loading can tell a scientist about its life feeding behaviour. Longer wings and low wing loading are typical of more pelagic species, while diving species have shorter wings. Species such as the wandering albatross , which forage over huge areas of sea, have
15561-536: The shore. But the origin was already forgotten, and the findings created the story that the coins was from the Spanish Armada of the 16th century. The wreck was rediscovered in 1972 by sport scuba divers Bengt-Olof Gustafsson ( Sweden ), Stefan Persson (Sweden), and Eystein Krohn-Dale (Norway). Under the bottom vegetation of the seabed, the sand bottom was littered with encrusted coins. The following year,
15694-553: The site was investigated by Bergen Sjøfartsmuseum. Little remained of the ship but more than 500 kilograms (1,100 lb) of gold and silver was salvaged. In total about 57,000 coins were found; 6,600 of them being gold coins—mostly the rare 1724 Dutch Gold Ducat minted in Utrecht with only a handful known prior to this find, and the rest silver coins. Parts of the treasure are kept at Bergen Sjøfartsmuseum in Bergen and also at
15827-416: The skewed sex ratio of western gulls in southern California. 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. Oil pollution in particular threatens species with restricted ranges or already depressed populations. Climate change mainly affect seabirds via changes to their habitat : various processes in
15960-432: 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. It may be that plunge divers are restricted in their hunting grounds to clear waters that afford a view of their prey from the air. While they are the dominant guild in the tropics, the link between plunge diving and water clarity
16093-457: The soil that increase the symbiotic efficiency. Predator partitioning occurs when species are attacked differently by different predators (or natural enemies more generally). For example, trees could differentiate their niche if they are consumed by different species of specialist herbivores , such as herbivorous insects. If a species density declines, so too will the density of its natural enemies, giving it an advantage. Thus, if each species
16226-444: The species, environmental variation..., and interactions between the two—in particular the abilities of some species, especially our own, to modify their environments and alter the range dynamics of many other species." Alteration of an ecological niche by its inhabitants is the topic of niche construction . The majority of species exist in a standard ecological niche, sharing behaviors, adaptations, and functional traits similar to
16359-496: The spread of the northern fulmar through the United Kingdom is attributed in part to the availability of discards. 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. Fisheries also have negative effects on seabirds, and these effects, particularly on the long-lived and slow-breeding albatrosses, are
16492-695: The squid eaten are too large to have been caught alive, and include mid-water species likely to be beyond the reach of albatrosses. Some species will also feed on other seabirds; for example, gulls, skuas and pelicans will often take eggs, chicks and even small adult seabirds from nesting colonies, while the giant petrels can kill prey up to the size of small penguins and seal pups. Seabirds' life histories are dramatically different from those of land birds. In general, they are K-selected , live much longer (anywhere between twenty and sixty years), delay breeding for longer (for up to ten years), and invest more effort into fewer young. Most species will only have one clutch
16625-495: The theme. 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
16758-491: The three species ( Red and Red-necked ) are oceanic for nine months of the year, crossing the equator to feed pelagically. Loons and grebes , which nest on lakes but winter at sea, are usually categorized as water birds, not seabirds. Although there are a number of sea ducks in the family Anatidae that are truly marine in the winter, by convention they are usually excluded from the seabird grouping. Many waders (or shorebirds) and herons are also highly marine, living on
16891-499: The tropicbirds and some penguins), but most of the colour in seabirds appears in the bills and legs. The plumage of seabirds is thought in many cases to be for camouflage , both defensive (the colour of US Navy battleships is the same as that of Antarctic prions , and in both cases it reduces visibility at sea) and aggressive (the white underside possessed by many seabirds helps hide them from prey below). The usually black wing tips help prevent wear, as they contain melanins that help
17024-809: The two species interact, how they use their resources, and the type of ecosystem in which they exist, among other factors. In addition, several mathematical models exist to quantify niche breadth, competition, and coexistence (Bastolla et al. 2005). However, regardless of methods used, niches and competition can be distinctly difficult to measure quantitatively, and this makes detection and demonstration of niche differentiation difficult and complex. Over time, two competing species can either coexist, through niche differentiation or other means, or compete until one species becomes locally extinct . Several theories exist for how niche differentiation arises or evolves given these two possible outcomes. Niche differentiation can arise from current competition. For instance, species X has
17157-639: The underbrush and escapes from its predators by shuffling from underbrush to underbrush. Its 'niche' is defined by the felicitous complementing of the thrasher's behavior and physical traits (camouflaging color, short wings, strong legs) with this habitat. Grinnellian niches can be defined by non-interactive (abiotic) variables and environmental conditions on broad scales. Variables of interest in this niche class include average temperature, precipitation, solar radiation, and terrain aspect which have become increasingly accessible across spatial scales. Most literature has focused on Grinnellian niche constructs, often from
17290-442: The underlying processes that affect Lotka-Volterra relationships within an ecosystem. The framework centers around "consumer-resource models" which largely split a given ecosystem into resources (e.g. sunlight or available water in soil) and consumers (e.g. any living thing, including plants and animals), and attempts to define the scope of possible relationships that could exist between the two groups. In contemporary niche theory,
17423-584: Was 22 metres (72 ft); another study, this time on Cory's shearwaters nesting near Corsica , found that of nine out of 61 male chicks that returned to breed at their natal colony bred in the burrow they were raised in, and two actually bred with their own mother. Colonies are usually situated on islands, cliffs or headlands, which land mammals have difficulty accessing. This is thought to provide protection to seabirds, which are often very clumsy on land. Coloniality often arises in types of bird that do not defend feeding territories (such as swifts , which have
17556-679: 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 ), 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. Island restoration techniques, pioneered by New Zealand, enable
17689-459: Was founded. Seabird Seabirds (also known as marine birds ) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution , as the same environmental problems and feeding niches have resulted in similar adaptations. The first seabirds evolved in the Cretaceous period , and modern seabird families emerged in
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