HOME

TheInfoList



OR:

The
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Procellariidae is a group of
seabird Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
s that comprises the fulmarine petrels, the
gadfly petrel The gadfly petrels or ''Pterodroma'' are a genus of about 35 species of petrels, part of the seabird order Procellariiformes. The gadfly petrels are named for their speedy weaving flight, as if evading gadflies (horse-fly, horseflies). The flight ...
s, the diving petrels, the prions, and the shearwaters. This family is part of the bird order
Procellariiformes Procellariiformes is an order (biology), order of seabirds that comprises four family (biology), families: the albatrosses, the Procellariidae, petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still call ...
(or tubenoses), which also includes the
albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Paci ...
es and the storm petrels. The procellariids are the most numerous family of tubenoses, and the most diverse. They range in size from the giant petrels with a wingspan of around , that are almost as large as the albatrosses, to the diving petrels with a wingspan of around that are similar in size to the little auks or dovekies in the family
Alcidae Auks or alcids are birds of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The family contains 25 extant or recently extinct species that are divided into ...
. Male and female birds are identical in appearance. The plumage color is generally dull, with blacks, whites, browns and grays. The birds feed on fish,
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
and
crustacea Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
, with many also taking fisheries discards and
carrion Carrion (), also known as a carcass, is the decaying flesh of dead animals. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
. Whilst agile swimmers and excellent in water, petrels have weak legs and can only shuffle on land, with the giant petrels of the genus '' Macronectes'' being the only two species that are capable of proper
terrestrial locomotion Terrestrial locomotion has evolution, evolved as animals adapted from ecoregion#Marine, aquatic to ecoregion#Terrestrial, terrestrial environments. Animal locomotion, Locomotion on land raises different problems than that in water, with reduced f ...
. All species are accomplished long-distance foragers, and many undertake long trans-equatorial migrations. They are colonial breeders, exhibiting long-term mate fidelity and site philopatry. In all species, a single white egg is laid each breeding season. The parents take it in turns to incubate the egg and to forage for food. The feeding area can be at a great distance from the nest site. The incubation times and chick-rearing periods are exceptionally long compared to other birds. Many procellariids have breeding populations of over several million pairs; others number fewer than 200 birds. Humans have traditionally exploited several species of
fulmar The fulmars are tube-nosed seabirds in the family Procellariidae. The family includes two extant species, and two extinct fossil species from the Miocene. Fulmars superficially resemble gulls, but are readily distinguished by their flight on s ...
and shearwater (known as muttonbirds) for food, fuel, and bait, a practice that continues in a controlled fashion today. Several species are threatened by
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived ther ...
attacking adults and chicks in breeding colonies and by long-line fisheries.


Taxonomy and evolution

The
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Procellariidae was introduced (as Procellaridæ) by the English zoologist
William Elford Leach William Elford Leach (2 February 1791 – 25 August 1836) was an English zoologist and marine biologist. Life and work Elford Leach was born at Hoe Gate, Plymouth, the son of an attorney. At the age of twelve he began a medical apprenticesh ...
in a guide to the contents of the
British Museum The British Museum is a Museum, public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is the largest in the world. It documents the story of human cu ...
published in 1820. The name is derived from the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
'' Procellaria'' which in turn is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word ''procella'' meaning "storm" or "gale". The type genus was named in 1758 by the Swedish naturalist
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in the tenth edition of his ''
Systema Naturae ' (originally in Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the syste ...
''. Procellariidae is one of families that make up the order
Procellariiformes Procellariiformes is an order (biology), order of seabirds that comprises four family (biology), families: the albatrosses, the Procellariidae, petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still call ...
. Before the introduction of
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
s, the traditional arrangement was to divide the Procellariiformes into a set of four families:
Diomedeidae Albatrosses, of the biological family (biology), family Diomedeidae, are large seabirds related to the procellariidae, procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the So ...
containing the albatrosses, Hydrobatidae containing all the storm petrels, Pelecanoididae containing the diving petrels and Procellariidae containing the petrels, shearwaters and fulmars. The family Hydrobatidae was further divided into two subfamilies, the northern storm petrels in Hydrobatinae and the southern or austral storm petrels in Oceanitinae. A 1998 analysis of mitochondrial cytochrome ''b'' sequences found there was deep genetic divergence between the two subfamilies. Subsequent large-scale multigene studies found that the two subfamilies were not
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
. The storm petrels were therefore split into two families: Hydrobatidae containing the northern storm petrels and Oceanitidae, containing the southern storm petrels. The multigene genetic studies found that the diving petrels in the family Pelecanoididae were nested within the family Procellariidae. As a result, the diving petrels was merged into Procellariidae. The molecular evidence suggests that the albatrosses were the first to diverge from the ancestral stock, and the austral storm petrels next, with the procellariids and northern storm petrels splitting most recently. Within the procellariid family, a genetic analysis based on the cytochrome ''b'' gene published in 2004 indicated that the genus ''Puffinus'' contained two distinct
clades In biology, a clade (), also known as a monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach to taxonomy ...
and was
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
. The genus was therefore split and a group of species moved into the resurrected genus '' Ardenna''. The other genera within the family were found to be
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
but the relationships between the genera remained unclear. This changed when a multigene genetic study published in 2021 provided a genus-level phylogeny of the family. There are 99
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of procellariid in 16
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
. The family has usually been broken up into four fairly distinct groups; the fulmarine petrels, the
gadfly petrel The gadfly petrels or ''Pterodroma'' are a genus of about 35 species of petrels, part of the seabird order Procellariiformes. The gadfly petrels are named for their speedy weaving flight, as if evading gadflies (horse-fly, horseflies). The flight ...
s, the prions, and the shearwaters. With the inclusion of the diving petrels there are now five main groups. * The fulmarine petrels include the largest procellariids, the giant petrels, as well as the two
fulmar The fulmars are tube-nosed seabirds in the family Procellariidae. The family includes two extant species, and two extinct fossil species from the Miocene. Fulmars superficially resemble gulls, but are readily distinguished by their flight on s ...
species, the
snow petrel The snow petrel (''Pagodroma nivea'') is the only member of the genus ''Pagodroma.'' It is one of only three birds that have been seen at the Geographic South Pole, along with the Antarctic petrel and the south polar skua, which has the most s ...
, the Antarctic petrel, and the Cape petrel. The fulmarine petrels are a diverse group with differing habits and appearances, but are linked morphologically by their
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
features, particularly the long prominent nasal tubes. *The four diving petrels are the smallest procellariids with lengths of around and wingspans of . They are compact birds with short wings that are adapted for use under water. They have a characteristic whirring flight and dive into the water without settling. They probably remain all year in the seas near their breeding sites. * The gadfly petrels, so named due to their helter-skelter flight, are the 35 species in the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Pterodroma''. The species vary from small to medium sizes, in length, and are long winged with short hooked bills. They are most closely related to Kerguelen petrel which is placed in its own genus ''Aphrodroma''. * The prions comprise seven species of true prion in the genus ''
Pachyptila ''Pachyptila'' is a genus of seabirds in the family (biology), family Procellariidae and the order (biology), order Procellariiformes. The members of this genus and the blue petrel form a sub-group called Prion (bird), prions. They range througho ...
'' and the closely related blue petrel. Often known in the past as whalebirds, three species have large bills filled with
lamellae Lamella (: lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * Lame ...
that they use to filter
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
somewhat as
baleen whale Baleen whales (), also known as whalebone whales, are marine mammals of the order (biology), parvorder Mysticeti in the infraorder Cetacea (whales, dolphins and porpoises), which use baleen plates (or "whalebone") in their mouths to sieve plankt ...
s do, though the old name derives from their association with whales, not their bills (though "prions" does, deriving from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
for "
saw A saw is a tool consisting of a tough blade, Wire saw, wire, or Chainsaw, chain with a hard toothed edge used to cut through material. Various terms are used to describe toothed and abrasive saws. Saws began as serrated materials, and when man ...
"). They are small procellariids, in length, with a prominent dark M-shaped mark across the upperwing of their grey plumage. All are restricted to the southern hemisphere. * The shearwaters are adapted for diving after prey instead of foraging on the ocean's surface; several species have been recorded diving deeper than . They are known for the long trans-equatorial migrations undertaken by many species. The shearwaters include the 20 or so species of the genus '' Puffinus'', seven species in the genus '' Ardenna'', as well as the five large '' Procellaria'' species and the four '' Calonectris'' species. While all these four genera are often known collectively as shearwaters, the ''Procellaria'' are called petrels in their common names.


Morphology and flight

The procellariids are small- to medium-sized seabirds. The largest, the southern giant petrel with a wingspan of , is almost as large as albatrosses; the smallest, the diving petrels have a wingspan of and are similar in size to little auks or dovekies in the family
Alcidae Auks or alcids are birds of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The family contains 25 extant or recently extinct species that are divided into ...
. There are no obvious differences between the sexes, although females tend to be slighter. Like all Procellariiformes, the procellariids have a characteristic tubular nasal passage used for olfaction. This ability to smell helps to locate patchily distributed prey at sea and may help locate nesting colonies. The
plumage Plumage () is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, there can b ...
of the procellariids is usually dull, with greys, bluish greys, blacks and browns being the usual colours, although some species have striking patterns such as the Cape petrel. The technique of
flight Flight or flying is the motion (physics), motion of an Physical object, object through an atmosphere, or through the vacuum of Outer space, space, without contacting any planetary surface. This can be achieved by generating aerodynamic lift ass ...
among procellariids depends on foraging methods. Compared to an average bird, all procellariids have a high
aspect ratio The aspect ratio of a geometry, geometric shape is the ratio of its sizes in different dimensions. For example, the aspect ratio of a rectangle is the ratio of its longer side to its shorter side—the ratio of width to height, when the rectangl ...
(meaning their wings are long and narrow) and a heavy
wing loading In aerodynamics, wing loading is the total weight of an aircraft or flying animal divided by the area of its wing. The stalling speed, takeoff speed and landing speed of an aircraft are partly determined by its wing loading. The faster an airc ...
. Therefore, they must maintain a high speed in order to remain in the air. Most procellariids use two techniques to do this, namely,
dynamic soaring Dynamic soaring is a flying technique used to gain energy by repeatedly crossing the boundary between air masses of different velocity. Such zones of wind gradient are generally found close to obstacles and close to the surface, so the technique is ...
and slope soaring. Dynamic soaring involves gliding across wave fronts, thus taking advantage of the vertical
wind gradient In common usage, wind gradient, more specifically wind speed gradient or wind velocity gradient, or alternatively shear wind, is the vertical component of the gradient of the mean horizontal wind speed in the lower atmosphere. It is the rate of ...
and minimising the effort required to stay in the air. Slope soaring is more straightforward: the procellariid turns to the wind, gaining height, from where it can then glide back down to the sea. Most procellariids aid their flight by means of flap-glides, where bursts of flapping are followed by a period of gliding; the amount of flapping dependent on the strength of the wind and the choppiness of the water. Because of the high speeds required for flight, procellariids need to either run or face into a strong wind in order to take off. The giant petrels share with the albatrosses an adaptation known as a shoulder-lock: a sheet of
tendon A tendon or sinew is a tough band of fibrous connective tissue, dense fibrous connective tissue that connects skeletal muscle, muscle to bone. It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tensi ...
that locks the wing when fully extended, allowing the wing to be kept up and out without any muscle effort. Gadfly petrels often feed on the wing, snapping prey without landing on the water. The flight of the smaller prions is similar to that of the storm petrels, being highly erratic and involving weaving and even looping the loop. The wings of all species are long and stiff. In some species of shearwater the wings are used to power the birds underwater while diving for prey. Their heavier wing loadings, in comparison with surface-feeding procellariids, allow these shearwaters to achieve considerable depths (below in the case of the
short-tailed shearwater The short-tailed shearwater or slender-billed shearwater (''Ardenna tenuirostris''; formerly ''Puffinus tenuirostris''), also called yolla or moonbird, and commonly known as the muttonbird in Australia, is the most abundant seabird species in A ...
). Procellariids generally have weak legs that are set back, and many species move around on land by resting on the breast and pushing themselves forward, often with the help of their wings. The exceptions to this are the two species of giant petrel, which have strong legs used when they feed on land.


Distribution and migration

The procellariids are present in all the world's oceans and most of the seas. They are absent from the
Bay of Bengal The Bay of Bengal is the northeastern part of the Indian Ocean. Geographically it is positioned between the Indian subcontinent and the Mainland Southeast Asia, Indochinese peninsula, located below the Bengal region. Many South Asian and Southe ...
and
Hudson Bay Hudson Bay, sometimes called Hudson's Bay (usually historically), is a large body of Saline water, saltwater in northeastern Canada with a surface area of . It is located north of Ontario, west of Quebec, northeast of Manitoba, and southeast o ...
, but are present year round or seasonally in the rest. The seas north of
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
are the centre of procellariid
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
, with the most species. Among the groups, the fulmarine petrels have a mostly polar distribution, with most species living around
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
and one, the northern fulmar ranging in the Northern
Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...
and
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
Oceans. Of the four species of diving petrel, two are found along the coasts of South America, while the remaining two have circumpolar distributions in the Southern Ocean. The prions are restricted to the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the world ocean, generally taken to be south of 60th parallel south, 60° S latitude and encircling Antarctica. With a size of , it is the seco ...
, and the gadfly petrels are found mostly in the tropics with some temperate species. The shearwaters are the most widespread group and breed in most temperate and tropical seas. Many procellariids undertake long annual migrations in the non-breeding season. Southern species of shearwater such as the
sooty shearwater The sooty shearwater (''Ardenna grisea'') is a medium-large shearwater in the seabird family Procellariidae. In New Zealand, it is also known by its Māori language, Māori name , and is harvested by Māori people for muttonbirding, muttonbird, l ...
and short-tailed shearwater, breeding on islands off
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, New Zealand and
Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
, undertake transequatorial migrations of millions of birds up to the waters off
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
and back each year during the austral winter.
Manx shearwater The Manx shearwater (''Puffinus puffinus'') is a medium-sized shearwater in the seabird family Procellariidae. The scientific name of this species records a name shift: Manx shearwaters were called Manks puffins in the 17th century. Puffin is an ...
s from the North Atlantic also undertake transequatorial migrations from Western Europe and North America to the waters off Brazil in the South Atlantic. The mechanisms of
navigation Navigation is a field of study that focuses on the process of monitoring and controlling the motion, movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navig ...
are poorly understood, but displacement experiments where individuals were removed from colonies and flown to far-flung release sites have shown that they are able to home in on their colonies with remarkable precision. A Manx shearwater released in
Boston Boston is the capital and most populous city in the Commonwealth (U.S. state), Commonwealth of Massachusetts in the United States. The city serves as the cultural and Financial centre, financial center of New England, a region of the Northeas ...
returned to its colony in Skomer,
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
within 13 days, a distance of 5,150 kilometres (3,200 mi).


Behaviour


Food and feeding

The diet of the procellariids is the most diverse of all the Procellariiformes, as are the methods employed to obtain it. With the exception of the giant petrels, all procellariids are exclusively marine, and the diet of all species is dominated by either fish,
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
,
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s and
carrion Carrion (), also known as a carcass, is the decaying flesh of dead animals. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
, or some combination thereof. The majority of species are surface feeders, obtaining food that has been pushed to the surface by other predators or currents, or have floated in death. Among the surface feeders some, principally the gadfly petrels, can obtain food by dipping from flight, while most of the rest feed while sitting on the water. These surface feeders are dependent on their prey being close to the surface, and for this reason procellariids are often found in association with other predators or oceanic convergences. Studies have shown strong associations between many different kinds of
seabird Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
s, including wedge-tailed shearwaters, and
dolphin A dolphin is an aquatic mammal in the cetacean clade Odontoceti (toothed whale). Dolphins belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontopori ...
s and
tuna A tuna (: tunas or tuna) is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae ( mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bul ...
, which push shoaling fish up towards the surface. The gadfly petrels and the Kerguelen petrel mainly feed at night. In so doing they can take advantage of the nocturnal migration of cephalopods and other food species towards the surface. The fulmarine petrels are generalists, which for the most part take many species of fish and crustacea. The giant petrels, uniquely for Procellariiformes, will feed on land, eating the carrion of other seabirds and seals. They will also attack the chicks of other seabirds. The diet of the giant petrels varies according to sex, with the females taking more
krill Krill ''(Euphausiids)'' (: krill) are small and exclusively marine crustaceans of the order (biology), order Euphausiacea, found in all of the world's oceans. The name "krill" comes from the Norwegian language, Norwegian word ', meaning "small ...
and the males more carrion. All the fulmarine petrels readily feed on fisheries discards at sea, a habit that has been implicated in (but not proved to have caused) the expansion in range of the northern fulmar in the Atlantic. The three larger prion species have bills filled with
lamellae Lamella (: lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * Lame ...
, which act as filters to sift
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
from the water. Water is forced through the lamellae and small prey items are collected. This technique is often used in conjunction with a method known as hydroplaning where the bird dips its bill beneath the surface and propels itself forward with wings and feet as if walking on the water. The diving petrels and many of the shearwaters are proficient divers. While it has long been known that they regularly dive from the surface to pursue prey, using their wings for propulsion, the depth that they are able to dive to was not appreciated (or anticipated) until scientists began to deploy maximum-depth recorders on foraging birds. Studies of both long-distance migrants such as the sooty shearwater and more sedentary species such as the black-vented shearwater have shown maximum diving depths of and . Tropical shearwaters, such as the wedge-tailed shearwater and the Sargasso shearwater, also dive in order to hunt, making the shearwaters the only tropical seabirds capable of exploiting that ecological niche (all other tropical seabirds feed close to the surface). Many other species of procellariid, from white-chinned petrels to
slender-billed prion The slender-billed prion (''Pachyptila belcheri'') or thin-billed prion, is a species of petrel, a seabird in the family Procellariidae. It is found in the southern oceans. Taxonomy The slender-billed prion was species description, formally ...
s, dive to a couple of metres below the surface, though not as proficiently or as frequently as the shearwaters.


Breeding


Colonies

The procellariids are colonial, nesting for the most part on islands. These colonies vary in size from over a million birds to just a few pairs, and can be densely concentrated or widely spaced. At one extreme the greater shearwater nests in concentrations of one pair per square metre in three colonies of more than one million pairs, whereas the giant petrels nest in clumped but widely spaced territories that barely qualify as colonial. Colonies are usually located near the coast, but some species nest far inland and even at high altitudes. Hutton's shearwater (''Puffinus huttoni'') breeds in burrows on the sea-facing mountainside of the Kaikoura Ranges on
South Island The South Island ( , 'the waters of Pounamu, Greenstone') is the largest of the three major islands of New Zealand by surface area, the others being the smaller but more populous North Island and Stewart Island. It is bordered to the north by ...
, New Zealand. The colonies are above sea level at a distance of from the coast. Other exceptions are Barau's petrel (''Pterodroma baraui'') that breeds at on the island of
Réunion Réunion (; ; ; known as before 1848) is an island in the Indian Ocean that is an overseas departments and regions of France, overseas department and region of France. Part of the Mascarene Islands, it is located approximately east of the isl ...
in the Indian Ocean, and the snow petrel (''Pagodroma nivea'') that breeds in Antarctica on mountain ledges up to from the open sea. Most seabirds are colonial, and the reasons for colonial behaviour are assumed to be similar, if incompletely understood by scientists. Procellariids for the most part have weak legs and are unable to easily take off, making them highly vulnerable to
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ian predators. Most procellariid colonies are located on islands that have historically been free of mammals; for this reason some species cannot help but be colonial as they are limited to a few locations to breed. Even species that breed on continental Antarctica, such as the Antarctic petrel, are forced by habitat preference (snow-free north-facing rock) to breed in just a few locations. Most procellariids' nests are in burrows or on the surface on open ground, with a smaller number nesting under the cover of vegetation (such as in a forest). All the fulmarine petrels bar the snow petrel nest in the open, the snow petrel instead nesting inside natural crevices. Of the rest of the procellariids the majority nest in burrows or crevices, with a few tropical species nesting in the open. There are several reasons for these differences. The fulmarine petrels are probably precluded from burrowing by their large size (the crevice-nesting snow petrel is the smallest fulmarine petrel) and the high latitudes they breed in, where frozen ground is difficult to burrow into. The smaller size of the other species, and their lack of agility on land, mean that even on islands free from mammal predators they are still vulnerable to skuas,
gull Gulls, or colloquially seagulls, are seabirds of the subfamily Larinae. They are most closely related to terns and skimmers, distantly related to auks, and even more distantly related to waders. Until the 21st century, most gulls were placed ...
s and other avian predators, something the aggressive oil-spitting fulmars are not. The chicks of all species are vulnerable to predation, but the chicks of fulmarine petrels can defend themselves in a similar fashion to their parents. In the higher latitudes there are thermal advantages to burrow nesting, as the temperature is more stable than on the surface, and there is no wind-chill to contend with. The absence of skuas, gulls and other predatory birds on
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
islands is why some shearwaters and two species of gadfly petrel ( Kermadec petrel and the herald petrel) can nest in the open. This has the advantages of reducing competition with burrow nesters from other species and allowing open-ground nesters to nest on
coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
line islets without soil for burrowing. Procellariids that burrow in order to avoid predation almost always attend their colonies nocturnally in order to reduce predation as well. Procellariids display high levels of
philopatry Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives ...
, exhibiting both natal philopatry and site fidelity. Natal philopatry, the tendency of a bird to breed close to where it hatched, is strong among all the Procellariiformes. The evidence for natal philopatry comes from several sources, not the least of which is the existence of several procellariid species that are endemic to a single island. The study of
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
provides evidence of restricted
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
between different colonies, and has been used to show philopatry in fairy prions.
Bird ringing Bird ringing (UK) or bird banding (US) is the attachment of a small, individually numbered metal or plastic tag to the leg or wing of a wild bird to enable individual identification. This helps in keeping track of the movements of the bird an ...
provides compelling evidence of philopatry; a study of Cory's shearwaters nesting near
Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...
found that nine out of 61 male chicks that returned to breed at their natal colony actually bred in the burrow they were raised in. This tendency towards philopatry is stronger in some species than others, and several species readily prospect potential new colony sites and colonise them. It is hypothesised that there is a cost to dispersing to a new site, the chance of not finding a mate of the same species, that selects against it for rarer species, whereas there is probably an advantage to dispersal for species that have colony sites that change dramatically during periods of glacial advance or retreat. There are differences in the tendency to disperse based on sex, with females being more likely to breed away from the natal site.


Mate and site fidelity

Procellariids, as well as having strong natal philopatry, exhibit strong site fidelity, returning to the same nesting site, burrow or territory in sequential years. The figure varies for different species but is high for most species, an estimated 91% for Bulwer's petrels. The strength of this fidelity can also vary with sex; almost 85% of male Cory's shearwaters return to the same burrow to breed the year after a successful breeding attempt, while the figure for females is around 76%. This tendency towards using the same site from year to year is matched by strong mate fidelity, with birds breeding with the same partner for many years; it has been suggested that the two are linked, with site fidelity acting as a means in which partnered birds could meet at the beginning of the breeding season. One pair of northern fulmars bred as a pair in the same site for 25 years. Like the
albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Paci ...
es the procellariids take several years to reach sexual maturity, though due to the greater variety of sizes and lifestyles, the age of first breeding stretches from two or three years in the smaller species to 12 years in the larger ones. The procellariids lack the elaborate breeding dances of the albatrosses, in no small part due to the tendency of most of them to attend colonies at night and breed in burrows, where visual displays are useless. The fulmarine petrels, which nest on the surface and attend their colonies diurnally, do use a repertoire of stereotyped behaviours such as cackling, preening, head waving and nibbling, but for most species courtship interactions are limited to some billing (rubbing the two bills together) in the burrow and the vocalisations made by all species. The calls serve a number of functions: they are used territorially to protect burrows or territories and to call for mates. Each call type is unique to a particular species and indeed it is possible for procellariids to identify the sex of the bird calling. It may also be possible to assess the quality of potential mates; a study of blue petrels found a link between the
rhythm Rhythm (from Greek , ''rhythmos'', "any regular recurring motion, symmetry") generally means a " movement marked by the regulated succession of strong and weak elements, or of opposite or different conditions". This general meaning of regular r ...
and duration of calls and the body mass of the bird. The ability of an individual to recognise its mate has been demonstrated in several species.


Breeding season

Like most seabirds, the majority of procellariids breed once a year. There are exceptions; many individuals of the larger species, such as the white-headed petrel, will skip a breeding season after successfully fledging a chick, and some of the smaller species, such as the Christmas shearwaters, breed on a nine-month schedule. Among those that breed annually, there is considerable variation as to the timing; some species breed in a fixed season while others breed all year round.
Climate Climate is the long-term weather pattern in a region, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteoro ...
and the availability of food resources are important influences on the timing of procellariid breeding; species that breed at higher
latitude In geography, latitude is a geographic coordinate system, geographic coordinate that specifies the north-south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from −90° at t ...
s always breed in the summer as conditions are too harsh in the winter. At lower latitudes many, but not all, species breed continuously. Some species breed seasonally to avoid competition with other species for burrows, to avoid predation or to take advantage of seasonally abundant food. Others, such as the tropical wedge-tailed shearwater, breed seasonally for unknown reasons. Among the species that exhibit seasonal breeding there can be high levels of synchronization, both of time of arrival at the colony and of lay date. Procellariids begin to attend their nesting colony around one month prior to laying. Males will arrive first and attend the colony more frequently than females, partly in order to protect a site or burrow from potential competitors. Prior to laying there is a period known as the pre-laying exodus in which both the male and female are away from the colony, building up reserves in order to lay and undertake the first incubation stint respectively. This pre-laying exodus can vary in length from 9 days (as in the Cape petrel) to around 50 days in Atlantic petrels. All procellariids lay a single white egg per pair per breeding season, in common with the rest of the Procellariiformes. The egg is large compared to that of other birds, weighing 6–24% of the female's weight. Immediately after laying the female goes back to sea to feed while the male takes over incubation. Incubation duties are shared by both sexes in shifts that vary in length between species, individuals and the stage of incubation. The longest recorded shift was 29 days by a Murphy's petrel from Henderson Island; the typical length of a
gadfly petrel The gadfly petrels or ''Pterodroma'' are a genus of about 35 species of petrels, part of the seabird order Procellariiformes. The gadfly petrels are named for their speedy weaving flight, as if evading gadflies (horse-fly, horseflies). The flight ...
stint is between 13 and 19 days. Fulmarine petrels, shearwaters and prions tend to have shorter stints, averaging between 3 and 13 days. Incubation takes a long time, from 40 days for the smaller species (such as prions) to around 55 days for the larger species. The incubation period is longer if eggs are abandoned temporarily; procellariid eggs are resistant to chilling and can still hatch after being left unattended for a few days. After hatching the chick is brooded by a parent until it is large enough to thermoregulate efficiently, and in some cases defend itself from predation. This guard stage lasts a short while for burrow-nesting species (2–3 days) but longer for surface nesting
fulmar The fulmars are tube-nosed seabirds in the family Procellariidae. The family includes two extant species, and two extinct fossil species from the Miocene. Fulmars superficially resemble gulls, but are readily distinguished by their flight on s ...
s (around 16–20 days) and giant petrels (20–30 days). After the guard stage both parents feed the chick. In many species the parent's foraging strategy alternates between short trips lasting 1–3 days and longer trips of 5 days. The shorter trips, which are taken over the continental shelf, benefit the chick with faster growth, but longer trips to more productive
pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
feeding grounds are needed for the parents to maintain their own body condition. The meals are composed of both prey items and stomach oil, an
energy Energy () is the physical quantity, quantitative physical property, property that is transferred to a physical body, body or to a physical system, recognizable in the performance of Work (thermodynamics), work and in the form of heat and l ...
-rich food that is lighter to carry than undigested prey items. This oil is created in a stomach organ known as a
proventriculus The proventriculus is part of the digestive system of birds.Encarta World English Dictionary orth American Edition(2007). ''Proventriculus''. Source: (accessed: December 18, 2007) An analogous organ exists in invertebrates and insects. Birds Th ...
from digested prey items, and gives procellariids and other Procellariiformes their distinctive musty smell. Chick development is quite slow for
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, with fledging taking place at around two months after hatching for the smaller species and four months for the largest species. The chicks of some species are abandoned by the parents; parents of other species continue to bring food to the nesting site after the chick has left. Chicks put on weight quickly and some can outweigh their parents, although they will slim down before they leave the nest. All procellariid chicks fledge by themselves, and there is no further parental care after fledging. Life expectancy of Procellariidae is between 15 and 20 years; the oldest recorded member was a northern fulmar that was over 50 years.


Relationship with humans


Exploitation

Procellariids have been a seasonally abundant source of food for people wherever people have been able to reach their colonies. Early records of human exploitation of shearwaters (along with albatrosses and cormorants) come from the remains of
hunter-gatherer A hunter-gatherer or forager is a human living in a community, or according to an ancestrally derived Lifestyle, lifestyle, in which most or all food is obtained by foraging, that is, by gathering food from local naturally occurring sources, esp ...
midden A midden is an old dump for domestic waste. It may consist of animal bones, human excrement, botanical material, mollusc shells, potsherds, lithics (especially debitage), and other artifacts and ecofacts associated with past human oc ...
s in southern Chile, where sooty shearwaters were taken 5000 years ago. More recently, procellariids have been hunted for food by Europeans, particularly the northern fulmar in Europe, and various species by
Inuit Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
, and sailors around the world. The hunting pressure on the Bermuda petrel, or cahow, was so intense that the species nearly became extinct and did go missing for 300 years. The name of one species, the providence petrel, is derived from its (seemingly) miraculous arrival on
Norfolk Island Norfolk Island ( , ; ) is an States and territories of Australia, external territory of Australia located in the Pacific Ocean between New Zealand and New Caledonia, directly east of Australia's Evans Head, New South Wales, Evans Head and a ...
, where it provided a windfall for starving European settlers; within ten years the providence petrel was extinct on Norfolk. Several species of procellariid have gone extinct in the Pacific since the arrival of humans, and their remains have been found in middens dated to that time. More sustainable shearwater harvesting industries developed in
Tasmania Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...
and New Zealand, where the practice of harvesting what are known as muttonbirds continues today.


Threats and conservation

While some species of procellariid have populations that number in the millions, many species are much less common and several are threatened with
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
. Human activities have caused dramatic declines in the numbers of some species, particularly species that were originally restricted to one island. According to the
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
43 species are listed as vulnerable or worse, with 12 critically endangered. Procellariids are threatened by many threats, but
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived ther ...
on their breeding grounds,
light pollution Light pollution is the presence of any unwanted, inappropriate, or excessive artificial Visible spectrum, lighting. In a descriptive sense, the term ''light pollution'' refers to the effects of any poorly implemented lighting sources, during the ...
, marine
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life or, more commonly, the site where such enterprise takes place ( a.k.a., fishing grounds). Commercial fisheries include wild fisheries and fish farm ...
particularly bycatch, pollution, exploitation and
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
are the main threats measures as the number of species involved. The most pressing threat for many species, particularly the smaller ones, comes from species introduced to their colonies. Procellariids overwhelmingly breed on islands away from land predators such as mammals, and for the most part have lost the defensive adaptations needed to deal with them (with the exception of the oil-spitting fulmarine petrels). The introduction of mammal predators such as
feral cat A feral cat or a stray cat is an unowned domestic cat (''Felis catus'') that lives outdoors and avoids human contact; it does not allow itself to be handled or touched, and usually remains hidden from humans. Feral cats may breed over dozens ...
s, rats,
mongoose A mongoose is a small terrestrial carnivorous mammal belonging to the family Herpestidae. This family has two subfamilies, the Herpestinae and the Mungotinae. The Herpestinae comprises 23 living species that are native to southern Europe, A ...
s and mice can have disastrous results for ecologically naïve seabirds. These predators can either directly attack and kill breeding adults, or, more commonly, attack eggs and chicks. Burrowing species that leave their young unattended at a very early stage are particularly vulnerable to attack. Studies on
grey-faced petrel The grey-faced petrel (''Pterodroma gouldi'') is a species of petrel endemic to the North Island of New Zealand. In New Zealand it is also known by its Māori name and (along with other species such as the sooty shearwater) as a muttonbird. ...
s breeding on New Zealand's Whale Island (Moutohora) have shown that a population under heavy pressure from Norway rats will produce virtually no young during a breeding season, whereas if the rats are controlled (through the use of poison), breeding success is much higher. That study highlighted the role that non-predatory introduced species can play in harming seabirds; introduced
rabbit Rabbits are small mammals in the family Leporidae (which also includes the hares), which is in the order Lagomorpha (which also includes pikas). They are familiar throughout the world as a small herbivore, a prey animal, a domesticated ...
s on the island caused little damage to the petrels, other than damaging their burrows, but they acted as a food source for the rats during the non-breeding season, which allowed rat numbers to be higher than they otherwise would be, resulting in more predators for the petrels to contend with. Interactions with introduced species can be quite complex. Gould's petrels breed only on two islands, Cabbage Tree Island and Boondelbah Island off Port Stephens (New South Wales). Introduced rabbits destroyed the forest understory on Cabbage Tree Island; this both increased the vulnerability of the petrels to natural predators and left them vulnerable to the sticky fruits of the birdlime tree (''
Pisonia umbellifera ''Ceodes umbellifera'', Synonym (taxonomy), synonym ''Pisonia umbellifera'', commonly known as the birdlime tree or bird catcher tree, is a species of plant in the Nyctaginaceae family. The evergreen shrub has soft wood, small pink or yellow flowe ...
''), a native plant. In the natural state these fruits lodge in the understory of the forest, but with the understory removed the fruits fall to the ground where the petrels move about, sticking to their feathers and making flight impossible. Larger species of procellariid face similar problems to the albatrosses with long-line fisheries. These species readily take offal from fishing boats and will steal bait from the long lines as they are being set, risking becoming snared on the hooks and drowning. In the case of the spectacled petrel this has led to the species undergoing a large decline and its listing as vulnerable. Diving species, most especially the shearwaters, are also vulnerable to
gillnet Gillnetting is a fishing method that uses gillnets: vertical panels of netting that hang from a line with regularly spaced floaters that hold the line on the surface of the water. The floats are sometimes called "corks" and the line with corks is ...
fisheries. Studies of gill-net fisheries show that shearwaters (sooty and short-tailed) compose 60% of the seabirds killed by gill-nets in Japanese waters and 40% in Monterey Bay, California in the 1980s, with the total number of shearwaters killed in Japan being between 65,000 and 125,000 per annum over the same study period (1978–1981). Procellariids are vulnerable to other threats as well. Ingestion of plastic flotsam is a problem for the family as it is for many other seabirds. Once swallowed, this plastic can cause a general decline in the fitness of the bird, or in some cases lodge in the gut and cause a blockage, leading to death by starvation. Procellariids are also vulnerable to general marine pollution, as well as oil spills. Some species, such as the Barau's petrel, the Newell's shearwater or the Cory's shearwater, which nest high up on large developed islands are victims of light pollution. Chicks that are fledging are attracted to streetlights and are unable to reach the sea. An estimated 20–40% of fledging Barau's petrels are attracted to the streetlights on Réunion. Conservationists are working with governments and fisheries to prevent further declines and increase populations of endangered procellariids. Progress has been made in protecting many colonies where most species are most vulnerable. On 20 June 2001, the Agreement on the Conservation of Albatrosses and Petrels was signed by seven major fishing nations. The agreement lays out a plan to manage fisheries by-catch, protect breeding sites, promote conservation in the industry, and research threatened species. The developing field of
island restoration The ecological restoration of islands, or island restoration, is the application of the principles of ecological restoration to islands and island groups. Islands, due to their isolation, are home to many of the world's endemic (ecology), endemic ...
, where introduced species are removed and native species and habitats restored, has been used in several procellariid recovery programmes. Invasive species such as rats, feral cats and pigs have been either removed or controlled in many remote islands in the tropical Pacific (such as the Northwestern Hawaiian Islands), around New Zealand (where island restoration was developed), and in the south Atlantic and
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
s. The grey-faced petrels of Whale Island (mentioned above) have been achieving much higher fledging successes after the introduced Norway rats were finally completely removed. At sea, procellariids threatened by long-line fisheries can be protected using 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 using bird scarers can all reduce the seabird by-catch. The Agreement on the Conservation of Albatrosses and Petrels came into force in 2004 and has been ratified by eight countries, Australia, Ecuador, New Zealand, Spain, South Africa, France, Peru and the United Kingdom. The treaty requires these countries to take specific actions to reduce by-catch and pollution and to remove introduced species from nesting islands.


See also

* List of Procellariidae species


References


Sources

* * * * *


External links


The Agreement for the Conservation of Albatrosses and Petrels (ACAP)
Treaty Website

: Don Roberson's family page {{Featured article Bird families Seabirds Extant Rupelian first appearances Taxa named by William Elford Leach