A bird colony is a large congregation of individuals of one or more species of bird that nest or roost in proximity at a particular location. Many kinds of birds are known to congregate in groups of varying size; a congregation of nesting birds is called a breeding colony. Colonial nesting birds include seabirds such as auks and albatrosses; wetland species such as herons; and a few passerines such as weaverbirds, certain blackbirds, and some swallows. A group of birds congregating for rest is called a communal roost. Evidence of colonial nesting has been found in non-neornithine birds (Enantiornithes), in sediments from the Late Cretaceous (Maastrichtian) of Romania. [1]
Approximately 13% of all bird species nest colonially. [2] Nesting colonies are very common among seabirds on cliffs and islands. Nearly 95% of seabirds are colonial, [3] leading to the usage, seabird colony, sometimes called a rookery. Many species of terns nest in colonies on the ground. Herons, egrets, storks, and other large waterfowl also nest communally in what are called heronries. Colony nesting may be an evolutionary response to a shortage of safe nesting sites and abundance or unpredictable food sources which are far away from the nest sites. [4] Colony-nesting birds often show synchrony in their breeding, meaning that chicks all hatch at once, with the implication that any predator coming along at that time would find more prey items than it could possibly eat. [2] [5]
What exactly constitutes a colony is a matter of definition. Tufted puffins, for example, are pelagic birds that nest on the steep slopes and rocky crevices on coastal cliffs, often on islands. Each pair excavates its own burrow. A congregation of puffin burrows on a marine island is considered a colony. [6] [7] [8] Sand martins (called bank swallows in North America) are seldom, if ever, observed to nest in solitude; such a dependence on social nesting would term the bird a colonial nester. [9] A more extreme example of colonial nesting is found in the weaverbird family. The sociable weaver of southern Africa constructs massive, multi-family dwellings of twigs and dry grasses, with many entrances leading to different nesting chambers, accommodating as many as a hundred nesting pairs. These structures resemble haystacks hanging from trees, and have been likened to apartment buildings or beehives. [10]
Some seabird colonies host thousands of nesting pairs of various species. Triangle Island, for example, the largest seabird colony in British Columbia, Canada, is home to auks, gulls, cormorants, shorebirds, and other birds, as well as some marine mammals. [7] Many seabirds show remarkable site fidelity, returning to the same burrow, nest or site for many years, and they will defend that site from rivals with great vigour. This increases breeding success, provides a place for returning mates to reunite, and reduces the costs of prospecting for a new site. Young adults breeding for the first time usually return to their natal colony, and often nest very close to where they hatched. Individual nesting sites at seabird colonies can be widely spaced, as in an albatross colony, or densely packed like an auk 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. Colony size is a major aspect of the social environment of colonial birds.
Some birds are known to nest alone when conditions are suitable, but not sometimes. The white-winged dove of southwestern North America was known to nest in large colonies when foraging areas could support such numbers. In 1978, in Tamaulipas, Mexico, researchers counted 22 breeding colonies of white-winged doves with a collective population size of more than eight million birds. But as habitat was transformed through urbanization or agriculture, the doves apparently spread out into smaller, less long-lived colonies. Today, these doves are observed to nest singlyin both urban and not urban areas. [11] [12]
The term colony has also been applied, perhaps misleadingly, to smaller nesting groups, such as forest-dwelling species that nest socially in a suitable stand of trees. The red-cockaded woodpecker, an endangered species of southeastern North America, is a social species that feeds and roosts in family groups, or clans. Clans nest and roost in clusters of tree cavities and use a cooperative breeding system. [13] Many parrot species are also extremely social. For example, the thick-billed parrot is another bird that nests and roosts communally; individuals of neighboring roosts has been observed to communicate with each other each morning to signal their readiness to form flocks for foraging. [14] However, these complex social structures in birds are a different sort of group behavior than what is normally considered colonial.
The habit of nesting in groups is believed to provide better survival against predators in several ways. Many colonies are situated in locations that are naturally free of predators. In other cases, the presence of many birds means there are more individuals available for defense. Also, synchronized breeding leads to such an abundance of offspring as to satiate predators.
For seabirds, colonies on islands have an obvious advantage over mainland colonies when it comes to protection from terrestrial predators. Other situations can also be found where bird colonies avoid predation. A study of yellow-rumped caciques in Peru found that the birds, which build enclosed, pouch-like nests in colonies of up to one hundred active nests, situate themselves near wasp nests, which provide some protection from tree-dwelling predators such as monkeys. [15] When other birds came to rob the nests, the caciques would cooperatively defend the colony by mobbing the invader. Mobbing, clearly a group effort, is well-known behavior, not limited to colonial species; the more birds participating in the mobbing, the more effective it is at driving off the predator. [2] Therefore, it has been theorized that the larger number of individuals available for vigilance and defense makes the colony a safer place for the individual birds nesting there. More pairs of eyes and ears are available to raise the alarm and rise to the occasion.
Another suggestion is that colonies act as information centers and birds that have not found good foraging sites are able to follow others, who have fared better, to find food. [16] This makes sense for foragers because the food source is one that can be locally abundant. This hypothesis would explain why the lesser kestrel, which feeds on insects, breeds in colonies, while the related common kestrel, which feeds on larger prey, does not. [17]
Colonial behaviour has its costs as well. It has been noted that parasitism by haematozoa is higher in colonial birds and it has been suggested that blood parasites might have shaped adaptations such as larger organs in the immune system and life-history traits. [18] Other costs include brood parasitism and competition for food and territory. [3] [4] Colony size is a factor in the ecological function of colony nesting. In a larger colony, increased competition for food can make it harder for parents to feed their chicks. [19]
The benefits and drawbacks for birds of nesting in groups seem to be highly situational. Although scientists have hypothesized about the advantages of group nesting in terms of enabling group defensive behavior, escape from predation by being surrounded by neighbors (called the selfish herd hypothesis), as well as escaping predators through sheer numbers, in reality, each of these functions evidently depends on a number of factors. [17] Clearly, there can be safety in numbers, but there is some doubt about whether it balances out against the tendency for conspicuous breeding colonies to attract predators, and some suggest that colonial breeding can actually make birds more vulnerable. [3] At a common tern colony in Minnesota, a study of spotted sandpipers observed to nest near the tern colony showed that the sandpipers that nested nearest the colony seemed to gain some protection from mammalian predators, but avian predators were apparently attracted to the colony and the sandpipers nesting there were actually more vulnerable. [20] In a study of a least tern colony in Connecticut, nocturnal avian predators in the form of black-crowned night herons and great horned owls were observed to repeatedly invade a colony, flying into the middle of the colony and meeting no resistance. [21]
For seabirds, the location of colonies on islands, which are inaccessible to terrestrial predators, is an obvious advantage. Islands where terrestrial predators have arrived in the form of rats, cats, foxes, etc., have devastated island seabird colonies. One well-studied case of this phenomenon has been the effect on common murre colonies on islands in Alaska, where foxes were introduced for fur farming. [22]
Colony-nesting birds have been used by humans as a source of food in the form of eggs and meat, down for bedding, feathers for quill pens, and guano for fertilizer. Over-exploitation can be devastating to a colony, or even to an entire population of a colonial species. For example, there was once a large seabird known as the great auk, which nested in colonies in the North Atlantic. Eggs and birds were used for a variety of purposes. Beginning in the 16th century, seafarers took the birds in especially great numbers to fill ships' larders, and by the mid-19th century, the great auk was extinct. Likewise, the short-tailed albatross of the North Pacific was heavily harvested at what seems to have been its primary colony on Torishima Island. Millions of birds were killed in less than two decades at the end of the 19th century. The species survives, though endangered. In North America, the extermination of the highly gregarious passenger pigeon has been well documented. The birds were hunted as if inexhaustible. Case in point: in 1871, in Wisconsin, an estimated 136 million pigeons nested in a dense congregation over a wide area; thousands of people were drawn to hunt the birds, shipping the squab to market by rail. The passenger pigeon is a famous example of a familiar bird going extinct in modern times. [23]
The use of seabird droppings as fertilizer, or guano, began with the Indigenous Peruvians, who collected it from sites along the coast of South America, such as the Chincha Islands. When, after the Spanish Conquest, the value of this fertilizer became known to the wider world, collection increased to the point where the supply nearly ran out, and other sources of guano had to be found. [24]
Seabird colonies can be predominately a single species or a mix of species:
Colonial nesting can be so close that multiple nests are interwoven:
Another variation is to have closely grouped separate nests:
Heronries and rookeries are often in the top of a stand of trees:
The tufted puffin, also known as crested puffin, is a relatively abundant medium-sized pelagic seabird in the auk family (Alcidae) found throughout the North Pacific Ocean. It is one of three species of puffin that make up the genus Fratercula and is easily recognizable by its thick red bill and yellow tufts.
Puffins are any of three species of small alcids (auks) in the bird genus Fratercula. These are pelagic seabirds that feed primarily by diving in the water. They breed in large colonies on coastal cliffs or offshore islands, nesting in crevices among rocks or in burrows in the soil. Two species, the tufted puffin and horned puffin, are found in the North Pacific Ocean, while the Atlantic puffin is found in the North Atlantic Ocean.
Seabirds are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same environmental problems and feeding niches have resulted in similar adaptations. The first seabirds evolved in the Cretaceous period, and modern seabird families emerged in the Paleogene.
The Atlantic puffin, also known as the common puffin, is a species of seabird in the auk family. It is the only puffin native to the Atlantic Ocean; two related species, the tufted puffin and the horned puffin being found in the northeastern Pacific. The Atlantic puffin breeds in Russia, Iceland, Ireland, Britain, Norway, Greenland, Newfoundland and Labrador, Nova Scotia, and the Faroe Islands, and as far south as Maine in the west and France in the east. It is most commonly found in the Westman Islands, Iceland. Although it has a large population and a wide range, the species has declined rapidly, at least in parts of its range, resulting in it being rated as vulnerable by the IUCN. On land, it has the typical upright stance of an auk. At sea, it swims on the surface and feeds on zooplankton, small fish, and crabs, which it catches by diving underwater, using its wings for propulsion.
Terns are seabirds in the family Laridae, subfamily Sterninae, that have a worldwide distribution and are normally found near the sea, rivers, or wetlands. Terns are treated in eleven genera in a subgroup of the family Laridae, which also includes several genera of gulls and the skimmers (Rynchops). They are slender, lightly built birds with long, forked tails, narrow wings, long bills, and relatively short legs. Most species are pale grey above and white below with a contrasting black cap to the head, but the marsh terns, the black-bellied tern, the Inca tern, and some noddies have dark body plumage for at least part of the year. The sexes are identical in appearance, but young birds are readily distinguishable from adults. Terns have a non-breeding plumage, which usually involves a white forehead and much-reduced black cap.
The skuas are a group of predatory seabirds with seven species forming the genus Stercorarius, the only genus in the family Stercorariidae. The three smaller skuas, the Arctic skua, the long-tailed skua, and the pomarine skua, are called jaegers in North American English.
The common tern is a seabird in the family Laridae. This bird has a circumpolar distribution, its four subspecies breeding in temperate and subarctic regions of Europe, Asia and North America. It is strongly migratory, wintering in coastal tropical and subtropical regions. Breeding adults have light grey upperparts, white to very light grey underparts, a black cap, orange-red legs, and a narrow pointed bill. Depending on the subspecies, the bill may be mostly red with a black tip or all black. There are several similar species, including the partly sympatric Arctic tern, which can be separated on plumage details, leg and bill colour, or vocalisations.
The great black-backed gull is the largest member of the gull family. Described by the Cornell Lab of Ornithology as "the king of the Atlantic waterfront", it is a very aggressive hunter, pirate, and scavenger. It breeds on the European and North American coasts and islands of the North Atlantic and is fairly sedentary, though some move farther south or inland to large lakes and reservoirs. The adult great black-backed gull has a white head, neck and underparts, dark grey wings and back, pink legs and yellow bill.
The pigeon guillemot is a species of bird in the auk family, Alcidae. One of three species in the genus Cepphus, it is most closely related to the spectacled guillemot. There are five subspecies of the pigeon guillemot; all subspecies, when in breeding plumage, are dark brown with a black iridescent sheen and a distinctive wing patch broken by a brown-black wedge. Its non-breeding plumage has mottled grey and black upperparts and white underparts. The long bill is black, as are the claws. The legs, feet, and inside of the mouth are red. It closely resembles the black guillemot, which is slightly smaller and lacks the dark wing wedge present in the pigeon guillemot.
The roseate tern is a species of tern in the family Laridae. The genus name Sterna is derived from Old English "stearn", "tern", and the specific dougallii refers to Scottish physician and collector Dr Peter McDougall (1777–1814). "Roseate" refers to the bird's pink breast in breeding plumage.
Forster's tern is a tern in the family Laridae. The genus name Sterna is derived from Old English "stearn", "tern", and forsteri commemorates the naturalist Johann Reinhold Forster.
The Aleutian tern is a migratory bird living in the subarctic region of the globe most of the year. It is frequently associated with the Arctic tern, which it closely resembles. While both species have a black cap, the Aleutian tern may be distinguished by its white forehead. During breeding season, the Arctic terns have bright red bills, feet, and legs while those of the Aleutian terns are black.
The greater crested tern, also called crested tern, swift tern, or great crested tern, is a tern in the family Laridae that nests in dense colonies on coastlines and islands in the tropical and subtropical Old World. Its five subspecies breed in the area from South Africa around the Indian Ocean to the central Pacific and Australia, all populations dispersing widely from the breeding range after nesting. This large tern is closely related to the royal and lesser crested terns, but can be distinguished by its size and bill colour.
The black noddy, also known as white-capped noddy, is a species of tern in the family Laridae. It is a medium-sized seabird with black plumage and a white cap that closely resembles the lesser noddy with which it was at one time considered conspecific. The black noddy has slightly darker plumage and dark rather than pale lores.
The great frigatebird is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific Ocean, such as Hawaii and the Galápagos Islands; in the Indian Ocean, colonies can be found in the Seychelles and Mauritius, and there is a tiny population in the South Atlantic, mostly on and around St. Helena and Boatswain Bird Island.
The least tern is a species of tern that breeds in North America and locally in northern South America. It is closely related to, and was formerly often considered conspecific with, the little tern of the Old World. Other close relatives include the yellow-billed tern and Peruvian tern, both from South America.
The fork-tailed storm petrel is a small seabird of the storm petrel family Hydrobatidae. It is the second-most abundant and widespread storm petrel and is the only bird in its family that is bluish-grey in colour.
The Antarctic tern is a seabird in the family Laridae. It ranges throughout the southern oceans and is found on small islands around Antarctica as well as on the shores of the mainland. Its diet consists primarily of small fish and crustaceans. It is very similar in appearance to the closely related Arctic tern, but it is stockier, and it is in its breeding plumage in the southern summer, when the Arctic tern has shed old feathers to get its non-breeding plumage. The Antarctic tern does not migrate like the Arctic tern does, but it can still be found on a very large range. This tern species is actually more closely related to the South American tern.
The term seabird is used for many families of birds in several orders that spend the majority of their lives at sea. Seabirds make up some, if not all, of the families in the following orders: Procellariiformes, Sphenisciformes, Pelecaniformes, and Charadriiformes. Many seabirds remain at sea for several consecutive years at a time, without ever seeing land. Breeding is the central purpose for seabirds to visit land. The breeding period is usually extremely protracted in many seabirds and may last over a year in some of the larger albatrosses; this is in stark contrast with passerine birds. Seabirds nest in single or mixed-species colonies of varying densities, mainly on offshore islands devoid of terrestrial predators. However, seabirds exhibit many unusual breeding behaviors during all stages of the reproductive cycle that are not extensively reported outside of the primary scientific literature.
The western cattle egret is a species of heron found in the tropics, subtropics and warm temperate zones. Formerly, most taxonomic authorities lumped this species and the eastern cattle egret together, but the two cattle egrets are now treated as separate species. Despite the similarities in plumage to the egrets of the genus Egretta, it is more closely related to the herons of Ardea. Originally native to parts of Asia, Africa and Europe, it has undergone a rapid expansion in its distribution and successfully colonised much of the rest of the world in the last century.