Procellariiformes
Encyclopedia
Procellariiformes is an order of seabird
s that comprises four families
: the albatross
es, petrels and shearwaters
, storm petrels, and diving petrel
s. Formerly called Tubinares and still called tubenoses in English, they are often referred to collectively as the petrel
s, a term that has been applied to all Procellariiformes or more commonly all the families except the albatrosses. They are almost exclusively pelagic (feeding in the open ocean). They have a cosmopolitan distribution
across the world's oceans, with the highest diversity being around New Zealand
.
Procellariiformes are colonial, mostly nesting on remote, predator-free islands. The larger species nest on the surface, while most smaller species nest in natural cavities and burrows
. They exhibit strong philopatry
, returning to their natal colony to breed and returning to the same nesting site over many years. Procellariiformes are monogamous
and form long-term pair bonds that are formed over several years and may last for the life of the pair. Only a single egg
is laid per nesting attempt, and usually only a single nesting attempt is made per year, although the larger albatrosses may only nest once every two years. Both parents participate in incubation
and chick rearing. Incubation times are long compared to other birds, as are fledgling periods. Once a chick has fledged there is no further parental care.
Procellariiformes have had a long relationship with humans. They have been important food sources for many people, and continue to be hunted as such in some parts of the world. They have also been the subject of numerous cultural depictions, particularly albatrosses. Procellariiformes are one of the most endangered bird taxa, with many species threatened with extinction
due to introduced predators
in their breeding colonies, marine pollution
and the danger of fisheries by-catch
. Scientists, conservationists, fishermen, and governments around the world are working to reduce the threats posed to them, and these efforts have led to the signing of the Agreement on the Conservation of Albatrosses and Petrels
, a legally binding international treaty signed in 2001.
word procella which means a violent wind or a storm, and iformes which is added to symbolize order. Therefore a violent wind or a storm refers to the fact that members of this order
like stormy and windy weather.
across the world's oceans and seas, although at the levels of family and genus there are some clear patterns. Antarctic Petrel
s, Thalassoica antarctica, have to fly over 100 mi (160.9 km) to get to the ocean from their breeding colonies in Antarctica, and Northern Fulmar
s breed on the northeastern tip of Greenland
, the furthest north piece of land. The most cosmopolitan family is the Procellariidae
, although within that family there are some gaps in distribution. The gadfly petrels, Pterodroma, have a generally tropical and temperate distribution, whereas the fulmarine petrel
s are mostly polar with some temperate species. The majority of the fulmarine petrels, along with the prions
, are confined to the southern hemisphere. The shearwater
s have the most widespread distribution, although they are absent from the Pacific north of Japan as breeding birds.
The storm petrels are almost as widespread as the procellariids, and fall into two distinct subfamilies; the Oceanitinae
have a mostly southern hemisphere distribution and the Hydrobatinae
are found mostly in the northern hemisphere. Amongst the albatross
es the majority of the family is restricted to the southern hemisphere, feeding and nesting in cool temperate areas, although one genus, Phoebastria, ranges across the north Pacific. The family is absent from the north Atlantic, although fossil records indicate they bred there once. Finally the diving-petrels are restricted to the southern hemisphere.
The various species within the order have a variety of migration
strategies. Some species undertake regular trans-equatorial migrations, such as the Sooty Shearwater
which annually migrates from its breeding grounds in New Zealand and Chile to the North Pacific off Japan
, Alaska
and California, an annual round trip of 64000 km (39,767.9 mi), the longest measured annual migration of any bird. A number of other petrel species undertake trans-equatorial migrations, including the Wilson's Storm Petrel and the Providence Petrel
, but no albatross
es cross do due to their reliance on wind assisted flight. There are other long-distant migrants within the order; Swinhoe's Storm Petrels breed in the western Pacific and migrates to the western Indian Ocean, and Bonin Petrel
s nesting in Hawaii
migrate to the coast of Japan during the non-breeding season.
Procellariiformes range in size from the very large Wandering Albatross
, at 11 kg (24.3 lb) and a 3.6 m (11.8 ft) wingspan, to the tiny Least Storm Petrel, at 20 g (0.705479242102239 oz) and a 32 cm (12.6 in) wingspan. They have their nostrils enclosed in one or two tubes on their straight, deeply grooved bills with hooked tips. The beaks are made up from several plates. Wings are long and narrow; feet are webbed, and the hind toe is undeveloped or non-existent. Plumage is predominantly black, white, and grey.
The order has a few unifying characteristics, starting with their tubular nasal passage which is used for olfaction
. This ability to smell helps to locate patchily distributed prey at sea and may also help locate their nests within nesting colonies. The structure of the bill
, which contains seven to nine distinct horny plates, is another unifying feature, although there are differences within the order. Petrels have a plate called the Maxillary unguis that forms a hook on the maxilla. The smaller members of the order have a comb-like mandible, made by the tomial plate, for plankton
feeding. Finally, they have a stomach oil stored in their proventriculus
that can be used as a food source during their long flights and also as a defense mechanism.
Procellariiformes have a need to lower their salt content due to their drinking of ocean water. All birds have an enlarged nasal gland at the base of the bill, above the eyes, which is inactive in species that don't require it. However, the Procellariiformes do require its use. Scientists are uncertain as to its exact processes, but do know that, in general terms, it removes salt from the system and forms a 5% saline solution that drips out of the nostrils, or is forcibly ejected in some petrels.
Most albatrosses and procellariids use two techniques minimise exertion while flying, namely, dynamic soaring
and slope soaring. The albatrosses and giant petrel
s share a morphological adaptation to aid in flight, a sheet of tendon
which locks the wing when fully extended, allowing the wing to be kept up and out without any muscle effort. Amongst the Oceanitinae
storm-petrels there are two unique flight patterns, one being surface pattering. In this they move across the water surface holding and moving their feet on the water's surface while holding steady above the water, and remaining stationary by hovering with rapid fluttering or by using the wind to anchor themselves in place. A similar flight method is thought to have been used by the extinct petrel family Diomedeoididae
. The White-faced Storm Petrel possesses a unique variation on pattering, holding its wings motionless and at an angle into the wind it pushes itself off the water's surface in a succession of bounding jumps.
Most are unable to walk well on land, and many species visit their remote breeding islands only at night. The exceptions are the huge albatrosses, several of the gadfly petrels and shearwaters and the fulmar-petrels. The latter can disable even large predatory birds with their obnoxious stomach oil
, which they can project some distance. This stomach oil is a digestive residue created in the foregut of all tubenoses except the diving petrels, and is used mainly for storage of energy rich food as well as for defence.
All Procellariiformes are colonial, predominantly breeding on offshore or oceanic islands. The few species that nest on continents do so in inhospitable environments such as dry deserts or on Antarctica. These colonies can vary from the widely spaced colonies of the giant petrel
s to the dense 3.6 million strong colonies of Leach's Storm Petrels. For almost all species the need to breed is the only reason that Procellariiformes return to land at all. Some of the larger petrels have to nest on windswept locations as they require wind to take off and forage for food. Within the colonies pairs defend usually small territories
(the giant petrels and some albatrosses can have very large territories) which is either the small area around the nest or a burrow. Competition between pairs can be intense, as can competition between species, particularly for burrows. Larger species of petrels will even kill the chicks and even adults of smaller species in disputes over burrows. Burrows and natural crevices are most commonly used by the smaller species; all the storm petrels and diving petrel
s are cavity nesters, as are many of the procellariids. The fulmarine petrel
s and some tropical gadfly petrel
s and shearwater
s are surface nesters, as are all the albatrosses. Colonies are often composed of several different species of both petrels and other seabirds.
Procellariiformes show high levels of philopatry
, both site fidelity and natal philopatry. Natal philopatry is the tendency of an individual bird to return to its natal colony to breed, often many years after leaving the colony as a chick. This tendency has been shown through ringing studies
and mitochondrial DNA
studies. In the ringing studies birds ringed as chicks are recapatured close to their original nests, a tendency which can be extreme at times; in Laysan Albatross
the average distance between hatching site and the site where a bird established its own territory was 22 m (72.2 ft), and a study of Cory's Shearwater
s nesting near Corsica
found that of nine out of 61 male chicks that returned to breed at their natal colony actually bred in the burrow they were raised in. Mitochondrial DNA
provides evidence of restricted gene flow
between different colonies, strongly suggesting philopatry.
The other type of philopatry exhibited is site fidelity, where pairs of birds return to the same nesting site for a number of years. Among the most extreme examples known of this tendency was the fidelity of a ringed
Northern Fulmar
that returned to the same nest site for 25 years. The average number of birds returning to the same nest sites is high in all species studied, with figures of around 91% for Bulwer's Petrel
s, and 85% of males and 76% of females for Cory's Shearwater
s (after a successful breeding attempt).
Procellariiformes are monogamous
breeders and form long term pair-bonds. These pair bonds take several years to develop in some species, particularly with the albatrosses. Having formed they will last for many breeding seasons, in some cases for the life of the pair. Petrel courtship can be an elaborate affair. It reaches its extreme with the albatrosses, where pairs of albatrosses spend many years perfecting and elaborate mating dances. These dances are composed of synchronised performances of various actions such as preening
, pointing, calling, bill clacking, staring, and combinations of such behaviours (like the sky-call). Each particular pair will develop their own individual version of the dance. The breeding behaviour of other Procellariiformes are less elaborate, although similar bonding behaviours are involved, particularly for the surface nesting procellariids. These can involve synchronised flights, mutual preening and calling
. Calls are important for helping birds locate potential mates and distinguish between species and may also serve a function in helping individuals assess the quality of potential mates. After pair formation has occurred calls also serve to help them reunite, the ability of individuals to recognise their own mate has also been demonstrated in several species.
Procellariiformes are k-selected. Breeding is delayed for several years after fledging
, sometimes for as long as ten years in the case of the largest species. Once they begin breeding, they make only a single breeding attempt per nesting season; even if the egg is lost early in the season, they will seldom re-lay. Large amounts of effort are placed into laying a single (proportionally) large egg
and raising a single chick. Procellariiformes are long-lived, the longest living albatross known survived for 51 years, but was probably older, even the tiny storm-petrels are known to have survived for 30 years.
, are able to nest on cycles slightly shorter than a year, and the large great albatross
es (genus Diomedea) nest in alternate years (if successful). Most temperate and polar species nest over the spring-summer, although some albatrosses and procellariids nest over the winter. In the tropics, some species can be found breeding throughout the year, but most nest in discreet periods. Procellariiformes return to nesting colonies as much as several months before laying, and attend their nest sites regularly before copulation. Prior to laying, females embark on a pre-laying exodus to build up energy reserves in order to lay the comparatively large egg.
When the female returns and lays, incubation is shared between the sexes, with the male taking the first incubation
stint and the female returning to sea. The duration of individual stints varies from just a few days to as much as several weeks, during which the incubating bird can lose a considerable amount of weight. The incubation period varies from species to species, around 40 days for the smallest storm-petrels but longer for the largest species; for albatrosses it can span 70 to 80 days, which is the longest incubation period of any bird.
Upon hatching, the chicks are semi-precocial
, having open eyes, a dense covering of white or grey down feather
s, and the ability to move around the nesting site. After hatching, the incubating adult remains with the chick for a number of days, a period known as the guard phase. In the case of most burrow-nesting species, this is only until the chick is able to thermoregulate
, usually two or three days. Diving-petrel chicks take longer to thermoregulate and have a longer guard phase than other burrow nesters. However, surface-nesting species, which have to deal with a greater range of weather and to contend with predators like skua
s and frigatebird
s, consequently have a longer guard phase (as long as two weeks in procellariids and three weeks in albatrosses).
The chick is fed by both parents. Chicks are fed on fish, squid, krill, and stomach oil
. Stomach oil is oil
composed of neutral dietary lipid
s that are the residue created by digestion
of the prey items. As an energy source for chicks it has several advantages over undigested prey, its calorific
value is around 9.6 kcal
per gram, which is only slightly lower than the value for diesel oil. This can be a real advantage for species that range over huge distances to provide food for hungry chicks. The oil is also used in defence. All Procellariiformes create stomach oil except the diving-petrels.
The chick fledge
s between two and nine months after hatching, almost twice as long as a gull of the same body mass. The reasons behind the length of time are associated with the distance from the breeding site to food. First, there are few predators at the nesting colonies, therefore there is no pressure to fledge quickly. Second, the time between feedings is long due to the distance from the nest site that adults forage, thus a chick that had a higher growth rate would stand a better chance of starving to death. The duration between feedings vary among species and during the stages of development. Small feeds are frequent during the guard phase, but afterward become less frequent.
's famous poem The Rime of the Ancient Mariner
, which in turn gave rise to the usage of albatross as metaphor for a burden
. There are few instances of petrel
s in culture, although there are sailors legends regarding the storm petrels, which are considered to warn of oncoming storms. In general petrels were considered to be "soul birds", representing the souls of drowned sailors, and it was considered unlucky to touch them. However, there also has been the belief that albatrosses were good omens and to kill one would bring bad luck.
In Russian, many petrel species from the Hydrobatidae and Pelecanoididae families of the order Procellariiformes are known as burevestnik
, which literally means 'the announcer of the storm'. When in 1901, the Russian writer Maxim Gorky
turned to the imagery of Subantarctic avifauna to describe Russian society's attitudes to the coming revolution, he used a storm-announcing petrel as the lead character of a poem that soon became popular in the revolutionary circles as "the battle anthem of the revolution". Although the species called "stormy petrel" in English is not one of those to which the burevestnik name is applied in Russian (it, in fact, is known in Russian as an entirely un-romantic kachurka), the English translators uniformly used the "stormy petrel" image in their translations of the poem, usually known in English as The Song of the Stormy Petrel
.
s and albatrosses along with those of other seabirds in 5,000 year old midden
s in Chile
, although it is likely that they were exploited prior to this. Since then many other marine cultures, both subsistence and industrial, have exploited Procellariiformes, in some cases almost to extinction
. Some cultures continue to harvest shearwaters (a practice known as muttonbirding
); for example the Māori of New Zealand
,who use a sustainable traditional method known as kaitiaki
tanga. In Alaska
, residents of Kodiak Island
harpoon Short-tailed Albatrosses, Diomedea albatrus, and until the late 1980s residents of Tristan Island in the Indian Ocean
have been harvesting the eggs of the Yellow-nosed Mollymawks, Diomedea chlororhynchos, and Sooty Albatrosses, Phoebetria fusca. Albatrosses and petrels are also now tourist draws in some locations, such as Taiaroa Head
. While such exploitation is non-consumptive, it can have deleterious effects that need careful management to protect both the birds and the tourism.
The albatrosses and petrels are "amongst the most severely threatened taxa worldwide". They face a variety of threats, the severity of which varies greatly from species to species. Several species are among the most common of seabirds, including the Wilson's Storm Petrel (an estimated 20 million individuals) and the Short-tailed Shearwater
(an estimated 30 million individuals); while the total population of some other species barely reaches more than two hundred individuals. There are less than 200 Magenta Petrel
s breeding on the Chatham Islands
, only 400 Zino's Petrel
s and only 80 Amsterdam Albatross
es. Only one species is thought to have become extinct since 1600, the Guadalupe Storm Petrel of Mexico
, although a number of species had died out before this. Numerous species are very poorly known, the Fiji Petrel
has only been seen a handful of times since its discovery and the breeding colonies of the New Zealand Storm Petrel, Hornby's Storm Petrel and Heinroth's Shearwater
have never been located. So little is known about the New Zealand Storm Petrel that is was thought extinct for 150 years until its rediscovery in 2003, although this record is dwarfed by that of the Bermuda Petrel
which was considered extinct for 330 years.
The principal threat to the albatrosses and larger species of procellariids is long-line fishing
. Bait set on hooks is attractive to foraging birds and many are hooked by the lines as they are set. As many as 100,000 albatrosses are hooked and drown each year on tuna
lines set out by long-line fisheries. However bad this number is, before 1991 and the ban on drift-net fisheries, it was estimated that 500,000 seabirds a year died as a result. This has led to spectacular declines in some species, as Procellariiformes are slow breeders and cannot replace their numbers fast enough.
Exotic species introduced
to the remote breeding colonies is also a threat to all types of Procellariiformes. These principally take the form of predators; most albatross and petrel species are clumsy on land and are unable to defend themselves from mammal
s such as rat
s, feral cat
s and pig
s. This phenomenon, known as ecological naivete
, has resulted in numerous declines in many species and has been strongly implicated in the extinction
of the Guadalupe Storm Petrel. Introduced herbivores can also cause problems if they unbalance the ecology
of the island; introduced rabbits destroyed the forest understory on Cabbage Tree Island off New South Wales
; this both increased the vulnerability of the Gould's Petrel
s nesting on the island to natural predators and left them vulnerable to the sticky fruits of the birdlime tree (Pisonia umbellifera
), 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.
In the past exploitation was a threat (see above), although this is less of a threat now. Other threats the ingestion of plastic flotsam. 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. This can also be picked up by foraging adults and fed to chicks, stunting their development and reducing the chances of successfully fledging. 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
and 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 and 45-60% of fledging Cory's Shearwater are attracted to the streetlights on Réunion
and Tenerife
, respectively.
.
In the Sibley-Ahlquist taxonomy
, the tubenoses are included in a greatly enlarged order "Ciconiiformes". This taxonomic
treatment is almost certainly erroneous, but the assumption of a close evolutionary relationship with other "higher waterbirds" – such as loon
s (Gaviiformes) and penguin
s (Sphenisciformes) – appears to be correct.
There are a total of around 125 living species of Procellariiformes worldwide, and the order is typically divided into four extant and one prehistorically extinct families:
The Hydrobatidae's two subfamilies, Oceanitinae and Hydrobatinae, are probably better treated as distinct families.
Primodroma, a fossil
procellariiform from the Early Eocene London Clay
of England
, may belong to the Hydrobatidae (perhaps specifically to the Oceanitinae) or maybe the Diomedeoididae.
A few rather fragmentary Late Cretaceous
and Paleogene
fossils have been occasionally allied with or even placed in the Procellariiformes. Marinavis, Neogaeornis
, Novacaesareala
, Torotix
and Tytthostonyx
seem to be "higher waterbirds" but cannot be reliably assigned to any of the modern lineages; rather, most of them appear to be still very close to the last common ancestor of Procellariiformes, cormorant
s, loons, pelican
s, penguins, and perhaps also grebe
s, flamingo
s, stork
s, tropicbird
s and wader
s. If they can be assigned to a modern order (which is highly doubtful), with the possible exception of Marinavis they would probably not be considered Procellariiformes. Eopuffinus and Manu
on the other hand are more likely members of the Procellariiformes; the former might be an ancestral petrel, the latter an ancient albatross. As regards Lonchodytes
(or rather its type species
L. estesi), it is the best candidate for the most ancient procellariiform known to date; it pre-dates the evolutionary radiation
that brought about the modern families and hence would occupy a basal position in the order. Parascaniornis on the other hand was formerly assigned to the Procellariiformes by some, but it is actually a hesperornithiform
synonymous with Baptornis
.
, Divers, before the extinction event. Fossil records are rare but 16 million year old fossils show that Albatross
es and Shearwater
s haven't changed much since then. It is believed that they evolved first in the Southern Hemisphere
, even though the majority of the fossils have been found in the Northern Hemisphere
. This is likely due to the fact that there is more land to find fossils in the north. DNA evidence has confirmed common ancestry for all Procellariiformes, however, the taxonomy
within the order is complex and fluctuating. The fossil record of the diving petrels goes back to the Miocene
, with a species from that family being described in 2007. The most numerous fossils from the Paleogene are those from the extinct family Diomedeoididae
, fossils of which have been found in Central Europe and Iran.
Seabird
Seabirds are birds that have 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...
s that comprises four families
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
: the albatross
Albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds allied to the procellariids, storm-petrels and diving-petrels in the order Procellariiformes . They range widely in the Southern Ocean and the North Pacific...
es, petrels and shearwaters
Procellariidae
The family Procellariidae is a group of seabirds that comprises the fulmarine petrels, the gadfly petrels, the prions, and the shearwaters. This family is part of the bird order Procellariiformes , which also includes the albatrosses, the storm-petrels, and the diving petrels.The procellariids are...
, storm petrels, and diving petrel
Diving petrel
The diving petrels are seabirds in the bird order Procellariiformes. There are four very similar species all in the family Pelecanoididae and genus Pelecanoides , distinguished only by small differences in the coloration of their plumage and their bill construction.Diving petrels are auk-like small...
s. Formerly called Tubinares and still called tubenoses in English, they are often referred to collectively as the petrel
Petrel
Petrels are tube-nosed seabirds in the bird order Procellariiformes. The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group...
s, a term that has been applied to all Procellariiformes or more commonly all the families except the albatrosses. They are almost exclusively pelagic (feeding in the open ocean). They have a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...
across the world's oceans, with the highest diversity being around New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...
.
Procellariiformes are colonial, mostly nesting on remote, predator-free islands. The larger species nest on the surface, while most smaller species nest in natural cavities and burrows
Burrows
Burrows is a provincial electoral division in the Canadian province of Manitoba. It was created by redistribution in 1957, and formally came into existence in the provincial election of 1958. The riding is located in the northern part of Winnipeg....
. They exhibit strong philopatry
Philopatry
Broadly, philopatry is the behaviour of remaining in, or returning to, an individual's birthplace. More specifically, in ecology philopatry is the behaviour of elder offspring sharing the parental burden in the upbringing of their siblings, a classic example of kin selection...
, returning to their natal colony to breed and returning to the same nesting site over many years. Procellariiformes are monogamous
Monogamy
Monogamy /Gr. μονός+γάμος - one+marriage/ a form of marriage in which an individual has only one spouse at any one time. In current usage monogamy often refers to having one sexual partner irrespective of marriage or reproduction...
and form long-term pair bonds that are formed over several years and may last for the life of the pair. Only a single egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
is laid per nesting attempt, and usually only a single nesting attempt is made per year, although the larger albatrosses may only nest once every two years. Both parents participate in incubation
Avian incubation
Incubation refers to the process by which certain oviparous animals hatch their eggs, and to the development of the embryo within the egg. The most vital factor of incubation is the constant temperature required for its development over a specific period. Especially in domestic fowl, the act of...
and chick rearing. Incubation times are long compared to other birds, as are fledgling periods. Once a chick has fledged there is no further parental care.
Procellariiformes have had a long relationship with humans. They have been important food sources for many people, and continue to be hunted as such in some parts of the world. They have also been the subject of numerous cultural depictions, particularly albatrosses. Procellariiformes are one of the most endangered bird taxa, with many species threatened with extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
due to introduced predators
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...
in their breeding colonies, marine pollution
Pollution
Pollution is the introduction of contaminants into a natural environment that causes instability, disorder, harm or discomfort to the ecosystem i.e. physical systems or living organisms. Pollution can take the form of chemical substances or energy, such as noise, heat or light...
and the danger of fisheries by-catch
By-catch
The term “bycatch” is usually used for fish caught unintentionally in a fishery while intending to catch other fish. It may however also indicate untargeted catch in other forms of animal harvesting or collecting...
. Scientists, conservationists, fishermen, and governments around the world are working to reduce the threats posed to them, and these efforts have led to the signing of the Agreement on the Conservation of Albatrosses and Petrels
Agreement on the Conservation of Albatrosses and Petrels
The Agreement on the Conservation of Albatrosses and Petrels is a legally binding international treaty signed in 2001.It was created in order to halt the drastic decline of seabird populations in the Southern Hemisphere, particularly albatrosses and procellariids...
, a legally binding international treaty signed in 2001.
Etymology
Procellariiformes comes from the LatinLatin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
word procella which means a violent wind or a storm, and iformes which is added to symbolize order. Therefore a violent wind or a storm refers to the fact that members of this order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
like stormy and windy weather.
Distribution and movements
The Procellariiformes have a cosmopolitan distributionCosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...
across the world's oceans and seas, although at the levels of family and genus there are some clear patterns. Antarctic Petrel
Antarctic Petrel
The Antarctic Petrel is a boldly marked dark brown and white petrel, found in Antarctica, most commonly in the Ross and Weddell seas. They eat Antarctic krill, fish, and small squid...
s, Thalassoica antarctica, have to fly over 100 mi (160.9 km) to get to the ocean from their breeding colonies in Antarctica, and Northern Fulmar
Northern Fulmar
The Northern Fulmar, Fulmarus glacialis, Fulmar, or Arctic Fulmar is a highly abundant sea bird found primarily in subarctic regions of the north Atlantic and north Pacific oceans. Fulmars come in one of two color morphs: a light one which is almost entirely white, and a dark one which is...
s breed on the northeastern tip of Greenland
Greenland
Greenland is an autonomous country within the Kingdom of Denmark, located between the Arctic and Atlantic Oceans, east of the Canadian Arctic Archipelago. Though physiographically a part of the continent of North America, Greenland has been politically and culturally associated with Europe for...
, the furthest north piece of land. The most cosmopolitan family is the Procellariidae
Procellariidae
The family Procellariidae is a group of seabirds that comprises the fulmarine petrels, the gadfly petrels, the prions, and the shearwaters. This family is part of the bird order Procellariiformes , which also includes the albatrosses, the storm-petrels, and the diving petrels.The procellariids are...
, although within that family there are some gaps in distribution. The gadfly petrels, Pterodroma, have a generally tropical and temperate distribution, whereas the fulmarine petrel
Fulmarine petrel
The fulmarine petrels or fulmar-petrels are a distinct group of petrels within the procellariidae family. They are the most variable of the four groups within the Procellariidae, differing greatly in size and biology. They do have, however, have a unifying feature, their skull, and in particular...
s are mostly polar with some temperate species. The majority of the fulmarine petrels, along with the prions
Prion (bird)
The Prions are small petrels in the genera Pachyptila and Halobaena. They form one of the four groups within the Procellariidae , along with the gadfly petrels, shearwaters and fulmarine petrels....
, are confined to the southern hemisphere. The shearwater
Shearwater
Shearwaters are medium-sized long-winged seabirds. There are more than 30 species of shearwaters, a few larger ones in the genus Calonectris and many smaller species in the genus Puffinus...
s have the most widespread distribution, although they are absent from the Pacific north of Japan as breeding birds.
The storm petrels are almost as widespread as the procellariids, and fall into two distinct subfamilies; the Oceanitinae
Oceanitinae
Oceanitinae is one of two subfamilies of storm-petrels. Because the distribution of the subfamily is predominantly in the southern hemisphere they are sometimes known as the southern storm-petrels. The subfamily contains five genera in which there are eight species...
have a mostly southern hemisphere distribution and the Hydrobatinae
Hydrobatinae
Hydrobatinae is one of two subfamilies of storm petrels. The distribution of the subfamily is predominantly in the northern hemisphere. The subfamily contains two genera in which there are 14 species...
are found mostly in the northern hemisphere. Amongst the albatross
Albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds allied to the procellariids, storm-petrels and diving-petrels in the order Procellariiformes . They range widely in the Southern Ocean and the North Pacific...
es the majority of the family is restricted to the southern hemisphere, feeding and nesting in cool temperate areas, although one genus, Phoebastria, ranges across the north Pacific. The family is absent from the north Atlantic, although fossil records indicate they bred there once. Finally the diving-petrels are restricted to the southern hemisphere.
The various species within the order have a variety of migration
Bird migration
Bird migration is the regular seasonal journey undertaken by many species of birds. Bird movements include those made in response to changes in food availability, habitat or weather. Sometimes, journeys are not termed "true migration" because they are irregular or in only one direction...
strategies. Some species undertake regular trans-equatorial migrations, such as the Sooty Shearwater
Sooty Shearwater
The Sooty Shearwater is a medium-large shearwater in the seabird family Procellariidae. In New Zealand it is also known by its Māori name tītī and as "muttonbird", like its relatives the Wedge-tailed Shearwater and the Australian Short-tailed Shearwater The Sooty Shearwater (Puffinus griseus) is...
which annually migrates from its breeding grounds in New Zealand and Chile to the North Pacific off Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
, Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
and California, an annual round trip of 64000 km (39,767.9 mi), the longest measured annual migration of any bird. A number of other petrel species undertake trans-equatorial migrations, including the Wilson's Storm Petrel and the Providence Petrel
Providence Petrel
The Providence Petrel is a species that burrows in one location; isolated Lord Howe Island, some 800km from the Australian mainland in the Tasman Sea....
, but no albatross
Albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds allied to the procellariids, storm-petrels and diving-petrels in the order Procellariiformes . They range widely in the Southern Ocean and the North Pacific...
es cross do due to their reliance on wind assisted flight. There are other long-distant migrants within the order; Swinhoe's Storm Petrels breed in the western Pacific and migrates to the western Indian Ocean, and Bonin Petrel
Bonin Petrel
The Bonin Petrel, Pterodroma hypoleuca, is a seabird in the family Procellariidae. It is a small gadfly petrel that lives in the waters of the north west Pacific and nests on islands south of Japan and in the Northwestern Hawaiian Islands...
s nesting in Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...
migrate to the coast of Japan during the non-breeding season.
Morphology and flight
Wandering Albatross
The Wandering Albatross, Snowy Albatross or White-winged Albatross, Diomedea exulans, is a large seabird from the family Diomedeidae, which has a circumpolar range in the Southern Ocean. It was the first species of albatross to be described, and was long considered the same species as the Tristan...
, at 11 kg (24.3 lb) and a 3.6 m (11.8 ft) wingspan, to the tiny Least Storm Petrel, at 20 g (0.705479242102239 oz) and a 32 cm (12.6 in) wingspan. They have their nostrils enclosed in one or two tubes on their straight, deeply grooved bills with hooked tips. The beaks are made up from several plates. Wings are long and narrow; feet are webbed, and the hind toe is undeveloped or non-existent. Plumage is predominantly black, white, and grey.
The order has a few unifying characteristics, starting with their tubular nasal passage which is used for olfaction
Olfaction
Olfaction is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates...
. This ability to smell helps to locate patchily distributed prey at sea and may also help locate their nests within nesting colonies. The structure of the bill
Beak
The beak, bill or rostrum is an external anatomical structure of birds which is used for eating and for grooming, manipulating objects, killing prey, fighting, probing for food, courtship and feeding young...
, which contains seven to nine distinct horny plates, is another unifying feature, although there are differences within the order. Petrels have a plate called the Maxillary unguis that forms a hook on the maxilla. The smaller members of the order have a comb-like mandible, made by the tomial plate, for plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
feeding. Finally, they have a stomach oil stored in their proventriculus
Proventriculus
The proventriculus is part of the digestive system of birds, invertebrates and insects.-Birds:The proventriculus is a standard part of avian anatomy...
that can be used as a food source during their long flights and also as a defense mechanism.
Procellariiformes have a need to lower their salt content due to their drinking of ocean water. All birds have an enlarged nasal gland at the base of the bill, above the eyes, which is inactive in species that don't require it. However, the Procellariiformes do require its use. Scientists are uncertain as to its exact processes, but do know that, in general terms, it removes salt from the system and forms a 5% saline solution that drips out of the nostrils, or is forcibly ejected in some petrels.
Dynamic soaring
Dynamic soaring is a flying technique used to gain energy by repeatedly crossing the boundary between air masses of significantly different velocity...
and slope soaring. The albatrosses and giant petrel
Giant petrel
Giant petrels is a genus, Macronectes, from the family Procellariidae and consist of two species. They are the largest birds from this family...
s share a morphological adaptation to aid in flight, a sheet of tendon
Tendon
A tendon is a tough band of fibrous connective tissue that usually connects muscle to bone and is capable of withstanding tension. Tendons are similar to ligaments and fasciae as they are all made of collagen except that ligaments join one bone to another bone, and fasciae connect muscles to other...
which locks the wing when fully extended, allowing the wing to be kept up and out without any muscle effort. Amongst the Oceanitinae
Oceanitinae
Oceanitinae is one of two subfamilies of storm-petrels. Because the distribution of the subfamily is predominantly in the southern hemisphere they are sometimes known as the southern storm-petrels. The subfamily contains five genera in which there are eight species...
storm-petrels there are two unique flight patterns, one being surface pattering. In this they move across the water surface holding and moving their feet on the water's surface while holding steady above the water, and remaining stationary by hovering with rapid fluttering or by using the wind to anchor themselves in place. A similar flight method is thought to have been used by the extinct petrel family Diomedeoididae
Diomedeoididae
The Diomedeoididae are a prehistoric family of seabirds. The family was in the order Procellariiformes which today is composed of the albatrosses and petrels. At present the family contains a single genus, Diomedeoides, in which there are three described species...
. The White-faced Storm Petrel possesses a unique variation on pattering, holding its wings motionless and at an angle into the wind it pushes itself off the water's surface in a succession of bounding jumps.
Most are unable to walk well on land, and many species visit their remote breeding islands only at night. The exceptions are the huge albatrosses, several of the gadfly petrels and shearwaters and the fulmar-petrels. The latter can disable even large predatory birds with their obnoxious stomach oil
Stomach oil
Stomach oil is the light oil composed of neutral dietary lipids found in the proventriculus of birds in the order Procellariiformes. All albatrosses, procellarids and storm-petrels use the oil...
, which they can project some distance. This stomach oil is a digestive residue created in the foregut of all tubenoses except the diving petrels, and is used mainly for storage of energy rich food as well as for defence.
Breeding colonies
Giant petrel
Giant petrels is a genus, Macronectes, from the family Procellariidae and consist of two species. They are the largest birds from this family...
s to the dense 3.6 million strong colonies of Leach's Storm Petrels. For almost all species the need to breed is the only reason that Procellariiformes return to land at all. Some of the larger petrels have to nest on windswept locations as they require wind to take off and forage for food. Within the colonies pairs defend usually small territories
Territory (animal)
In ethology the term territory refers to any sociographical area that an animal of a particular species consistently defends against conspecifics...
(the giant petrels and some albatrosses can have very large territories) which is either the small area around the nest or a burrow. Competition between pairs can be intense, as can competition between species, particularly for burrows. Larger species of petrels will even kill the chicks and even adults of smaller species in disputes over burrows. Burrows and natural crevices are most commonly used by the smaller species; all the storm petrels and diving petrel
Diving petrel
The diving petrels are seabirds in the bird order Procellariiformes. There are four very similar species all in the family Pelecanoididae and genus Pelecanoides , distinguished only by small differences in the coloration of their plumage and their bill construction.Diving petrels are auk-like small...
s are cavity nesters, as are many of the procellariids. The fulmarine petrel
Fulmarine petrel
The fulmarine petrels or fulmar-petrels are a distinct group of petrels within the procellariidae family. They are the most variable of the four groups within the Procellariidae, differing greatly in size and biology. They do have, however, have a unifying feature, their skull, and in particular...
s and some tropical gadfly petrel
Gadfly petrel
The gadfly petrels are seabirds in the bird order Procellariiformes. These medium to large petrels feed on food items picked from the ocean surface....
s and shearwater
Shearwater
Shearwaters are medium-sized long-winged seabirds. There are more than 30 species of shearwaters, a few larger ones in the genus Calonectris and many smaller species in the genus Puffinus...
s are surface nesters, as are all the albatrosses. Colonies are often composed of several different species of both petrels and other seabirds.
Procellariiformes show high levels of philopatry
Philopatry
Broadly, philopatry is the behaviour of remaining in, or returning to, an individual's birthplace. More specifically, in ecology philopatry is the behaviour of elder offspring sharing the parental burden in the upbringing of their siblings, a classic example of kin selection...
, both site fidelity and natal philopatry. Natal philopatry is the tendency of an individual bird to return to its natal colony to breed, often many years after leaving the colony as a chick. This tendency has been shown through ringing studies
Bird ringing
Bird ringing or bird banding is a technique used in the study of wild birds, by attaching a small, individually numbered, metal or plastic tag to their legs or wings, so that various aspects of the bird's life can be studied by the ability to re-find the same individual later...
and mitochondrial DNA
Mitochondrial DNA
Mitochondrial DNA is the DNA located in organelles called mitochondria, structures within eukaryotic cells that convert the chemical energy from food into a form that cells can use, adenosine triphosphate...
studies. In the ringing studies birds ringed as chicks are recapatured close to their original nests, a tendency which can be extreme at times; in Laysan Albatross
Laysan Albatross
The Laysan Albatross, Phoebastria immutabilis, is a large seabird that ranges across the North Pacific. This small two-tone gull-like albatross is the second most common seabird in the Hawaiian Islands, with an estimated population of 2.5 million birds, and is currently expanding its range to new...
the average distance between hatching site and the site where a bird established its own territory was 22 m (72.2 ft), and a study of Cory's Shearwater
Cory's Shearwater
The Cory's Shearwater is a large shearwater in the seabird family Procellariidae.This species breeds on islands and cliffs in the Mediterranean, with the odd outpost on the Atlantic coast of Iberia. The nest is on open ground or among rocks or less often in a burrow where one white egg is laid,...
s nesting near Corsica
Corsica
Corsica is an island in the Mediterranean Sea. It is located west of Italy, southeast of the French mainland, and north of the island of Sardinia....
found that of nine out of 61 male chicks that returned to breed at their natal colony actually bred in the burrow they were raised in. Mitochondrial DNA
Mitochondrial DNA
Mitochondrial DNA is the DNA located in organelles called mitochondria, structures within eukaryotic cells that convert the chemical energy from food into a form that cells can use, adenosine triphosphate...
provides evidence of restricted gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
between different colonies, strongly suggesting philopatry.
The other type of philopatry exhibited is site fidelity, where pairs of birds return to the same nesting site for a number of years. Among the most extreme examples known of this tendency was the fidelity of a ringed
Bird ringing
Bird ringing or bird banding is a technique used in the study of wild birds, by attaching a small, individually numbered, metal or plastic tag to their legs or wings, so that various aspects of the bird's life can be studied by the ability to re-find the same individual later...
Northern Fulmar
Northern Fulmar
The Northern Fulmar, Fulmarus glacialis, Fulmar, or Arctic Fulmar is a highly abundant sea bird found primarily in subarctic regions of the north Atlantic and north Pacific oceans. Fulmars come in one of two color morphs: a light one which is almost entirely white, and a dark one which is...
that returned to the same nest site for 25 years. The average number of birds returning to the same nest sites is high in all species studied, with figures of around 91% for Bulwer's Petrel
Bulwer's Petrel
The Bulwer's Petrel is a small petrel in the family Procellariidae, and is one of two species in the genus Bulweria . This bird is named after the Scottish naturalist James Bulwer.- Description :...
s, and 85% of males and 76% of females for Cory's Shearwater
Cory's Shearwater
The Cory's Shearwater is a large shearwater in the seabird family Procellariidae.This species breeds on islands and cliffs in the Mediterranean, with the odd outpost on the Atlantic coast of Iberia. The nest is on open ground or among rocks or less often in a burrow where one white egg is laid,...
s (after a successful breeding attempt).
Pair bonds and life history
Monogamy
Monogamy /Gr. μονός+γάμος - one+marriage/ a form of marriage in which an individual has only one spouse at any one time. In current usage monogamy often refers to having one sexual partner irrespective of marriage or reproduction...
breeders and form long term pair-bonds. These pair bonds take several years to develop in some species, particularly with the albatrosses. Having formed they will last for many breeding seasons, in some cases for the life of the pair. Petrel courtship can be an elaborate affair. It reaches its extreme with the albatrosses, where pairs of albatrosses spend many years perfecting and elaborate mating dances. These dances are composed of synchronised performances of various actions such as preening
Personal grooming
Personal grooming is the art of cleaning, grooming, and maintaining parts of the body. It is a species-typical behavior that is controlled by neural circuits in the brain.- In humans :...
, pointing, calling, bill clacking, staring, and combinations of such behaviours (like the sky-call). Each particular pair will develop their own individual version of the dance. The breeding behaviour of other Procellariiformes are less elaborate, although similar bonding behaviours are involved, particularly for the surface nesting procellariids. These can involve synchronised flights, mutual preening and calling
Bird song
Bird vocalization includes both bird calls and bird songs. In non-technical use, bird songs are the bird sounds that are melodious to the human ear. In ornithology and birding, songs are distinguished by function from calls.-Definition:The distinction between songs and calls is based upon...
. Calls are important for helping birds locate potential mates and distinguish between species and may also serve a function in helping individuals assess the quality of potential mates. After pair formation has occurred calls also serve to help them reunite, the ability of individuals to recognise their own mate has also been demonstrated in several species.
Procellariiformes are k-selected. Breeding is delayed for several years after fledging
Fledge
Fledge is the stage in a young bird's life when the feathers and wing muscles are sufficiently developed for flight. It also describes the act of a chick's parents raising it to a fully grown state...
, sometimes for as long as ten years in the case of the largest species. Once they begin breeding, they make only a single breeding attempt per nesting season; even if the egg is lost early in the season, they will seldom re-lay. Large amounts of effort are placed into laying a single (proportionally) large egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
and raising a single chick. Procellariiformes are long-lived, the longest living albatross known survived for 51 years, but was probably older, even the tiny storm-petrels are known to have survived for 30 years.
Nesting and chick rearing
The majority of Procellariiformes nest once a year and do so seasonally. Some tropical shearwaters, like the Christmas ShearwaterChristmas Shearwater
The Christmas Shearwater, Puffinus nativitatis, is a medium-sized shearwater of the tropical Central Pacific. It is a poorly known species due to its remote nesting habits, and it has not been extensively studied at sea either....
, are able to nest on cycles slightly shorter than a year, and the large great albatross
Great albatross
The great albatrosses are seabirds in the genus Diomedea in the albatross family. The genus Diomedea formerly included all albatrosses except the sooty albatrosses, but in 1996 the genus was split with the mollymawks and the North Pacific albatrosses both being elevated to separate genera...
es (genus Diomedea) nest in alternate years (if successful). Most temperate and polar species nest over the spring-summer, although some albatrosses and procellariids nest over the winter. In the tropics, some species can be found breeding throughout the year, but most nest in discreet periods. Procellariiformes return to nesting colonies as much as several months before laying, and attend their nest sites regularly before copulation. Prior to laying, females embark on a pre-laying exodus to build up energy reserves in order to lay the comparatively large egg.
When the female returns and lays, incubation is shared between the sexes, with the male taking the first incubation
Avian incubation
Incubation refers to the process by which certain oviparous animals hatch their eggs, and to the development of the embryo within the egg. The most vital factor of incubation is the constant temperature required for its development over a specific period. Especially in domestic fowl, the act of...
stint and the female returning to sea. The duration of individual stints varies from just a few days to as much as several weeks, during which the incubating bird can lose a considerable amount of weight. The incubation period varies from species to species, around 40 days for the smallest storm-petrels but longer for the largest species; for albatrosses it can span 70 to 80 days, which is the longest incubation period of any bird.
Upon hatching, the chicks are semi-precocial
Precocial
In biology, the term precocial refers to species in which the young are relatively mature and mobile from the moment of birth or hatching. The opposite developmental strategy is called "altricial," where the young are born or hatched helpless. Extremely precocial species may be called...
, having open eyes, a dense covering of white or grey down feather
Down feather
The down of birds is a layer of fine feathers found under the tougher exterior feathers. Very young birds are clad only in down. Powder down is a specialized type of down found only in a few groups of birds. Down is a fine thermal insulator and padding, used in goods such as jackets, bedding,...
s, and the ability to move around the nesting site. After hatching, the incubating adult remains with the chick for a number of days, a period known as the guard phase. In the case of most burrow-nesting species, this is only until the chick is able to thermoregulate
Thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different...
, usually two or three days. Diving-petrel chicks take longer to thermoregulate and have a longer guard phase than other burrow nesters. However, surface-nesting species, which have to deal with a greater range of weather and to contend with predators like skua
Skua
The skuas are a group of seabirds with about seven species forming the family Stercorariidae and the genus Stercorarius. The three smaller skuas are called jaegers in North America....
s and frigatebird
Frigatebird
The frigatebirds are a family, Fregatidae, of seabirds. There are five species in the single genus Fregata. They are also sometimes called Man of War birds or Pirate birds. Since they are related to the pelicans, the term "frigate pelican" is also a name applied to them...
s, consequently have a longer guard phase (as long as two weeks in procellariids and three weeks in albatrosses).
The chick is fed by both parents. Chicks are fed on fish, squid, krill, and stomach oil
Stomach oil
Stomach oil is the light oil composed of neutral dietary lipids found in the proventriculus of birds in the order Procellariiformes. All albatrosses, procellarids and storm-petrels use the oil...
. Stomach oil is oil
Oil
An oil is any substance that is liquid at ambient temperatures and does not mix with water but may mix with other oils and organic solvents. This general definition includes vegetable oils, volatile essential oils, petrochemical oils, and synthetic oils....
composed of neutral dietary lipid
Lipid
Lipids constitute a broad group of naturally occurring molecules that include fats, waxes, sterols, fat-soluble vitamins , monoglycerides, diglycerides, triglycerides, phospholipids, and others...
s that are the residue created by digestion
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....
of the prey items. As an energy source for chicks it has several advantages over undigested prey, its calorific
Calorie
The calorie is a pre-SI metric unit of energy. It was first defined by Nicolas Clément in 1824 as a unit of heat, entering French and English dictionaries between 1841 and 1867. In most fields its use is archaic, having been replaced by the SI unit of energy, the joule...
value is around 9.6 kcal
Calorie
The calorie is a pre-SI metric unit of energy. It was first defined by Nicolas Clément in 1824 as a unit of heat, entering French and English dictionaries between 1841 and 1867. In most fields its use is archaic, having been replaced by the SI unit of energy, the joule...
per gram, which is only slightly lower than the value for diesel oil. This can be a real advantage for species that range over huge distances to provide food for hungry chicks. The oil is also used in defence. All Procellariiformes create stomach oil except the diving-petrels.
The chick fledge
Fledge
Fledge is the stage in a young bird's life when the feathers and wing muscles are sufficiently developed for flight. It also describes the act of a chick's parents raising it to a fully grown state...
s between two and nine months after hatching, almost twice as long as a gull of the same body mass. The reasons behind the length of time are associated with the distance from the breeding site to food. First, there are few predators at the nesting colonies, therefore there is no pressure to fledge quickly. Second, the time between feedings is long due to the distance from the nest site that adults forage, thus a chick that had a higher growth rate would stand a better chance of starving to death. The duration between feedings vary among species and during the stages of development. Small feeds are frequent during the guard phase, but afterward become less frequent.
Role in culture
The most important family in terms of cultural importance is the albatrosses, which have been described by one author as "the most legendary of birds". Albatrosses have featured in poetry in the form of Samuel Taylor ColeridgeSamuel Taylor Coleridge
Samuel Taylor Coleridge was an English poet, Romantic, literary critic and philosopher who, with his friend William Wordsworth, was a founder of the Romantic Movement in England and a member of the Lake Poets. He is probably best known for his poems The Rime of the Ancient Mariner and Kubla...
's famous poem The Rime of the Ancient Mariner
The Rime of the Ancient Mariner
The Rime of the Ancient Mariner is the longest major poem by the English poet Samuel Taylor Coleridge, written in 1797–98 and was published in 1798 in the first edition of Lyrical Ballads. Modern editions use a later revised version printed in 1817 that featured a gloss...
, which in turn gave rise to the usage of albatross as metaphor for a burden
Albatross (metaphor)
The word 'albatross' is sometimes used metaphorically to mean a psychological burden that feels like a curse. It is an allusion to Samuel Taylor Coleridge's poem The Rime of the Ancient Mariner ....
. There are few instances of petrel
Petrel
Petrels are tube-nosed seabirds in the bird order Procellariiformes. The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group...
s in culture, although there are sailors legends regarding the storm petrels, which are considered to warn of oncoming storms. In general petrels were considered to be "soul birds", representing the souls of drowned sailors, and it was considered unlucky to touch them. However, there also has been the belief that albatrosses were good omens and to kill one would bring bad luck.
In Russian, many petrel species from the Hydrobatidae and Pelecanoididae families of the order Procellariiformes are known as burevestnik
Burevestnik
Burevestnik was the All-Union Voluntary Sports Society of students and teachers of the most part of high schools and universities in the USSR, established in 1957 .Since 1959 the society was a member of the International University Sports Federation...
, which literally means 'the announcer of the storm'. When in 1901, the Russian writer Maxim Gorky
Maxim Gorky
Alexei Maximovich Peshkov , primarily known as Maxim Gorky , was a Russian and Soviet author, a founder of the Socialist Realism literary method and a political activist.-Early years:...
turned to the imagery of Subantarctic avifauna to describe Russian society's attitudes to the coming revolution, he used a storm-announcing petrel as the lead character of a poem that soon became popular in the revolutionary circles as "the battle anthem of the revolution". Although the species called "stormy petrel" in English is not one of those to which the burevestnik name is applied in Russian (it, in fact, is known in Russian as an entirely un-romantic kachurka), the English translators uniformly used the "stormy petrel" image in their translations of the poem, usually known in English as The Song of the Stormy Petrel
The Song of the Stormy Petrel
"The Song of the Stormy Petrel" is a short piece of revolutionary literature written by the Russian writer Maxim Gorky in 1901. Written in a variation of unrhymed trochaic tetrameter with occasional Pyrrhic substitutions, it is considered poetry.-History:...
.
Exploitation
Albatrosses and petrels have been important food sources for humans for as long as people have been able to reach their remote breeding colonies. Amongst the earliest known examples of this is the remains of shearwaterShearwater
Shearwaters are medium-sized long-winged seabirds. There are more than 30 species of shearwaters, a few larger ones in the genus Calonectris and many smaller species in the genus Puffinus...
s and albatrosses along with those of other seabirds in 5,000 year old midden
Midden
A midden, is an old dump for domestic waste which may consist of animal bone, human excrement, botanical material, vermin, shells, sherds, lithics , and other artifacts and ecofacts associated with past human occupation...
s in Chile
Chile
Chile ,officially the Republic of Chile , is a country in South America occupying a long, narrow coastal strip between the Andes mountains to the east and the Pacific Ocean to the west. It borders Peru to the north, Bolivia to the northeast, Argentina to the east, and the Drake Passage in the far...
, although it is likely that they were exploited prior to this. Since then many other marine cultures, both subsistence and industrial, have exploited Procellariiformes, in some cases almost to extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
. Some cultures continue to harvest shearwaters (a practice known as muttonbirding
Muttonbirding
Muttonbirding is a seasonal harvesting activity, which may be recreational or commercial, of the chicks of petrels, especially shearwater species, for food, oil and feathers...
); for example the Māori of New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...
,who use a sustainable traditional method known as kaitiaki
Kaitiaki
Kaitiaki is a New Zealand term used for the Māori concept of guardianship, for the sky, the sea, and the land. A kaitiaki is a guardian, and the process and practices of protecting and looking after the environment are referred to as kaitiakitanga and include rāhui and tapuThe term kaitiaki is also...
tanga. In Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
, residents of Kodiak Island
Kodiak Island
Kodiak Island is a large island on the south coast of the U.S. state of Alaska, separated from the Alaska mainland by the Shelikof Strait. The largest island in the Kodiak Archipelago, Kodiak Island is the second largest island in the United States and the 80th largest island in the world, with an...
harpoon Short-tailed Albatrosses, Diomedea albatrus, and until the late 1980s residents of Tristan Island in the Indian Ocean
Indian Ocean
The Indian Ocean is the third largest of the world's oceanic divisions, covering approximately 20% of the water on the Earth's surface. It is bounded on the north by the Indian Subcontinent and Arabian Peninsula ; on the west by eastern Africa; on the east by Indochina, the Sunda Islands, and...
have been harvesting the eggs of the Yellow-nosed Mollymawks, Diomedea chlororhynchos, and Sooty Albatrosses, Phoebetria fusca. Albatrosses and petrels are also now tourist draws in some locations, such as Taiaroa Head
Taiaroa Head
Taiaroa Head is a headland at the end of the Otago Peninsula in New Zealand, overlooking the mouth of the Otago Harbour. It lies within the city limits of Dunedin...
. While such exploitation is non-consumptive, it can have deleterious effects that need careful management to protect both the birds and the tourism.
Threats and conservation
See Also: Introduced mammals on seabird breeding islandsIntroduced mammals on seabird breeding islands
Seabirds include some of the most threatened taxa anywhere in the world. For example, of extant albatross species, 82% are listed as threatened, endangered, or critically endangered by the International Union for Conservation of Nature. The two leading threats to seabirds are accidental bycatch by...
Short-tailed Shearwater
The Short-tailed Shearwater or Slender-billed Shearwater , also called Yolla or Moonbird, and commonly known as the muttonbird in Australia, is the most abundant seabird species in Australian waters, and is one of the few Australian native birds in which the chicks are commercially harvested...
(an estimated 30 million individuals); while the total population of some other species barely reaches more than two hundred individuals. There are less than 200 Magenta Petrel
Magenta Petrel
The Magenta Petrel or Chatham Island Taiko is a small seabird in the gadfly petrel genus, Pterodroma....
s breeding on the Chatham Islands
Chatham Islands
The Chatham Islands are an archipelago and New Zealand territory in the Pacific Ocean consisting of about ten islands within a radius, the largest of which are Chatham Island and Pitt Island. Their name in the indigenous language, Moriori, means Misty Sun...
, only 400 Zino's Petrel
Zino's Petrel
The Zino's Petrel or Freira, Pterodroma madeira, is a small seabird in the gadfly petrel genus which is endemic to the island of Madeira. This long-winged petrel has a grey back and wings, with a dark "W" marking across the wings, and a grey upper tail...
s and only 80 Amsterdam Albatross
Amsterdam Albatross
The Amsterdam Albatross or Amsterdam Island Albatross, Diomedea amsterdamensis, is a huge albatross which breeds only on Amsterdam Island in the southern Indian Ocean. It was only described in 1983, and was thought by some researchers to be a sub-species of the Wandering Albatross, exulans...
es. Only one species is thought to have become extinct since 1600, the Guadalupe Storm Petrel of Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
, although a number of species had died out before this. Numerous species are very poorly known, the Fiji Petrel
Fiji Petrel
The Fiji Petrel , also known as MacGillivray's Petrel, is a small, dark gadfly petrel.The Fiji Petrel was originally known from one immature specimen found in 1855 on Gau Island, Fiji by naturalist John MacGillivray on board 'HMS Herald' who took the carcass to the British Museum in London...
has only been seen a handful of times since its discovery and the breeding colonies of the New Zealand Storm Petrel, Hornby's Storm Petrel and Heinroth's Shearwater
Heinroth's Shearwater
Heinroth's Shearwater is a poorly known seabird in the family Procellariidae. Probably a close relative of the Little Shearwater or Audubon's Shearwater , it is distinguished by a long and slender bill and a brown-washed underside.This species is restricted to the seas around the Bismarck...
have never been located. So little is known about the New Zealand Storm Petrel that is was thought extinct for 150 years until its rediscovery in 2003, although this record is dwarfed by that of the Bermuda Petrel
Bermuda Petrel
The Bermuda Petrel, Pterodroma cahow, is a gadfly petrel. Commonly known in Bermuda as the Cahow, a name derived from its eerie cries, this nocturnal ground-nesting seabird is the national bird of Bermuda, and a symbol of hope for nature conservation. It was thought extinct for 330 years...
which was considered extinct for 330 years.
Long-line fishing
Longline fishing is a commercial fishing technique. It uses a long line, called the main line, with baited hooks attached at intervals by means of branch lines called "snoods". A snood is a short length of line, attached to the main line using a clip or swivel, with the hook at the other end....
. Bait set on hooks is attractive to foraging birds and many are hooked by the lines as they are set. As many as 100,000 albatrosses are hooked and drown each year on tuna
Tuna
Tuna is a salt water fish from the family Scombridae, mostly in the genus Thunnus. Tuna are fast swimmers, and some species are capable of speeds of . Unlike most fish, which have white flesh, the muscle tissue of tuna ranges from pink to dark red. The red coloration derives from myoglobin, an...
lines set out by long-line fisheries. However bad this number is, before 1991 and the ban on drift-net fisheries, it was estimated that 500,000 seabirds a year died as a result. This has led to spectacular declines in some species, as Procellariiformes are slow breeders and cannot replace their numbers fast enough.
Exotic species introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...
to the remote breeding colonies is also a threat to all types of Procellariiformes. These principally take the form of predators; most albatross and petrel species are clumsy on land and are unable to defend themselves from mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s such as rat
Rat
Rats are various medium-sized, long-tailed rodents of the superfamily Muroidea. "True rats" are members of the genus Rattus, the most important of which to humans are the black rat, Rattus rattus, and the brown rat, Rattus norvegicus...
s, feral cat
Feral cat
A feral cat is a descendant of a domesticated cat that has returned to the wild. It is distinguished from a stray cat, which is a pet cat that has been lost or abandoned, while feral cats are born in the wild; the offspring of a stray cat can be considered feral if born in the wild.In many parts of...
s and pig
Pig
A pig is any of the animals in the genus Sus, within the Suidae family of even-toed ungulates. Pigs include the domestic pig, its ancestor the wild boar, and several other wild relatives...
s. This phenomenon, known as ecological naivete
Island tameness
Island tameness is the tendency of many populations and species of animals living on isolated islands to lose their wariness of potential predators, particularly of large animals. The term is partly synonymous with ecological naïvete, which also has a wider meaning referring to the loss of...
, has resulted in numerous declines in many species and has been strongly implicated in the extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
of the Guadalupe Storm Petrel. Introduced herbivores can also cause problems if they unbalance the ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...
of the island; introduced rabbits destroyed the forest understory on Cabbage Tree Island off New South Wales
New South Wales
New South Wales is a state of :Australia, located in the east of the country. It is bordered by Queensland, Victoria and South Australia to the north, south and west respectively. To the east, the state is bordered by the Tasman Sea, which forms part of the Pacific Ocean. New South Wales...
; this both increased the vulnerability of the Gould's Petrel
Gould's Petrel
Gould's Petrel is a species of seabird in the Procellariidae family. It is a small petrel, 30 cm long with a wingspan of about 70 cm. It is largely grey above and white below with a blackish crown and hindneck and a black M-shaped band across the wings and rump.The subspecies P. l...
s nesting on the island to natural predators and left them vulnerable to the sticky fruits of the birdlime tree (Pisonia umbellifera
Pisonia umbellifera
Pisonia umbellifera, commonly known as the Bird-lime Tree, is a species of plant in the Nyctaginaceae family. It grows throughout the tropical Indo-Pacific. It is native to the Andaman Islands, Indonesia, Malaysia, the Philippines, Thailand, Vietnam, China, Taiwan, Hawaii and Madagascar and the...
), 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.
In the past exploitation was a threat (see above), although this is less of a threat now. Other threats the ingestion of plastic flotsam. 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. This can also be picked up by foraging adults and fed to chicks, stunting their development and reducing the chances of successfully fledging. Procellariids are also vulnerable to general marine pollution, as well as oil spills. Some species, such as the Barau's Petrel
Barau's Petrel
Barau's Petrel, Pterodroma baraui is a medium sized gadfly petrel from the family Procellariidae. Its main breeding site is the island of Réunion in the Indian Ocean.-Etymology:...
, the Newell's Shearwater
Newell's Shearwater
Newell's Shearwater or Hawaiian Shearwater is a seabird belonging to the genus Puffinus in the family Procellariidae. It belongs to a confusing group of shearwaters which are difficult to identify and whose classification is controversial...
and the Cory's Shearwater
Cory's Shearwater
The Cory's Shearwater is a large shearwater in the seabird family Procellariidae.This species breeds on islands and cliffs in the Mediterranean, with the odd outpost on the Atlantic coast of Iberia. The nest is on open ground or among rocks or less often in a burrow where one white egg is laid,...
, which nest high up on large developed islands are victims of light pollution. Chicks that are fledging
Fledge
Fledge is the stage in a young bird's life when the feathers and wing muscles are sufficiently developed for flight. It also describes the act of a chick's parents raising it to a fully grown state...
are attracted to streetlights and are unable to reach the sea. An estimated 20–40% of fledging Barau's Petrels and 45-60% of fledging Cory's Shearwater are attracted to the streetlights on Réunion
Réunion
Réunion is a French island with a population of about 800,000 located in the Indian Ocean, east of Madagascar, about south west of Mauritius, the nearest island.Administratively, Réunion is one of the overseas departments of France...
and Tenerife
Tenerife
Tenerife is the largest and most populous island of the seven Canary Islands, it is also the most populated island of Spain, with a land area of 2,034.38 km² and 906,854 inhabitants, 43% of the total population of the Canary Islands. About five million tourists visit Tenerife each year, the...
, respectively.
Taxonomy and systematics
At one point (until the beginning of the 20th century), the family Hydrobatidae was named Procellariidae, and the family now called Procellariidae was rendered "Puffinidae." The order itself was called Tubinares. A major early work on this group is F. DuCane Godman's Monograph of the Petrels, five fascicles, 1907—1910., with portraits of figures by John Gerrard KeulemansJohn Gerrard Keulemans
Johannes Gerardus Keulemans was a Dutch bird illustrator.-Biography and Work:...
.
In the Sibley-Ahlquist taxonomy
Sibley-Ahlquist taxonomy
The Sibley-Ahlquist taxonomy is a bird taxonomy proposed by Charles Sibley and Jon Edward Ahlquist. It is based on DNA-DNA hybridization studies conducted in the late 1970s and throughout the 1980s....
, the tubenoses are included in a greatly enlarged order "Ciconiiformes". This taxonomic
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
treatment is almost certainly erroneous, but the assumption of a close evolutionary relationship with other "higher waterbirds" – such as loon
Loon
The loons or divers are a group of aquatic birds found in many parts of North America and northern Eurasia...
s (Gaviiformes) and penguin
Penguin
Penguins are a group of aquatic, flightless birds living almost exclusively in the southern hemisphere, especially in Antarctica. Highly adapted for life in the water, penguins have countershaded dark and white plumage, and their wings have become flippers...
s (Sphenisciformes) – appears to be correct.
There are a total of around 125 living species of Procellariiformes worldwide, and the order is typically divided into four extant and one prehistorically extinct families:
- Family †DiomedeoididaeDiomedeoididaeThe Diomedeoididae are a prehistoric family of seabirds. The family was in the order Procellariiformes which today is composed of the albatrosses and petrels. At present the family contains a single genus, Diomedeoides, in which there are three described species...
(Early Oligocene – Early Miocene) - Family ProcellariidaeProcellariidaeThe family Procellariidae is a group of seabirds that comprises the fulmarine petrels, the gadfly petrels, the prions, and the shearwaters. This family is part of the bird order Procellariiformes , which also includes the albatrosses, the storm-petrels, and the diving petrels.The procellariids are...
(shearwaterShearwaterShearwaters are medium-sized long-winged seabirds. There are more than 30 species of shearwaters, a few larger ones in the genus Calonectris and many smaller species in the genus Puffinus...
s, fulmarine petrelFulmarine petrelThe fulmarine petrels or fulmar-petrels are a distinct group of petrels within the procellariidae family. They are the most variable of the four groups within the Procellariidae, differing greatly in size and biology. They do have, however, have a unifying feature, their skull, and in particular...
s, gadfly petrelGadfly petrelThe gadfly petrels are seabirds in the bird order Procellariiformes. These medium to large petrels feed on food items picked from the ocean surface....
s, and prionPrion (bird)The Prions are small petrels in the genera Pachyptila and Halobaena. They form one of the four groups within the Procellariidae , along with the gadfly petrels, shearwaters and fulmarine petrels....
s) - Family Diomedeidae (albatrossAlbatrossAlbatrosses, of the biological family Diomedeidae, are large seabirds allied to the procellariids, storm-petrels and diving-petrels in the order Procellariiformes . They range widely in the Southern Ocean and the North Pacific...
es) - Family Hydrobatidae (storm petrels)
- Family Pelecanoididae (diving petrelDiving petrelThe diving petrels are seabirds in the bird order Procellariiformes. There are four very similar species all in the family Pelecanoididae and genus Pelecanoides , distinguished only by small differences in the coloration of their plumage and their bill construction.Diving petrels are auk-like small...
s)
The Hydrobatidae's two subfamilies, Oceanitinae and Hydrobatinae, are probably better treated as distinct families.
Primodroma, a fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
procellariiform from the Early Eocene London Clay
London Clay
The London Clay Formation is a marine geological formation of Ypresian age which crops out in the southeast of England. The London Clay is well known for the fossils it contains. The fossils from the Lower Eocene indicate a moderately warm climate, the flora being tropical or subtropical...
of England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...
, may belong to the Hydrobatidae (perhaps specifically to the Oceanitinae) or maybe the Diomedeoididae.
A few rather fragmentary Late Cretaceous
Late Cretaceous
The Late Cretaceous is the younger of two epochs into which the Cretaceous period is divided in the geologic timescale. Rock strata from this epoch form the Upper Cretaceous series...
and Paleogene
Paleogene
The Paleogene is a geologic period and system that began 65.5 ± 0.3 and ended 23.03 ± 0.05 million years ago and comprises the first part of the Cenozoic Era...
fossils have been occasionally allied with or even placed in the Procellariiformes. Marinavis, Neogaeornis
Neogaeornis
Neogaeornis is a controversial prehistoric genus of diving bird. The single known species, Neogaeornis wetzeli, was described from fossils found in the Campanian to Maastrichtian Quiriquina Formation of Chile. It lived about 70-67 million years ago...
, Novacaesareala
Novacaesareala
Novacaesareala is a genus of prehistoric bird. It is known only from the fossil remains of a single partial wing of the species Novacaesareala hungerfordi...
, Torotix
Torotix
Torotix is a Late Cretaceous genus of aquatic bird. It lived along the shores of the Western Interior Seaway, but it is not clear whether it was a seabird or a freshwater bird, as it is only known from a humerus. Consequently, the genus is monotypic, with the single species Torotix clemensi.It...
and Tytthostonyx
Tytthostonyx
Tytthostonyx is a genus of prehistoric seabird. Found in the much-debated Hornerstown Formation which straddles the Cretaceous-Paleogene boundary 65 million years ago, this animal was apparently closely related to the ancestor of some modern birds, such as Procellariiformes and/or "Pelecaniformes"...
seem to be "higher waterbirds" but cannot be reliably assigned to any of the modern lineages; rather, most of them appear to be still very close to the last common ancestor of Procellariiformes, cormorant
Cormorant
The bird family Phalacrocoracidae is represented by some 40 species of cormorants and shags. Several different classifications of the family have been proposed recently, and the number of genera is disputed.- Names :...
s, loons, pelican
Pelican
A pelican, derived from the Greek word πελεκυς pelekys is a large water bird with a large throat pouch, belonging to the bird family Pelecanidae....
s, penguins, and perhaps also grebe
Grebe
A grebe is a member of the Podicipediformes order, a widely distributed order of freshwater diving birds, some of which visit the sea when migrating and in winter...
s, flamingo
Flamingo
Flamingos or flamingoes are gregarious wading birds in the genus Phoenicopterus , the only genus in the family Phoenicopteridae...
s, stork
Stork
Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family Ciconiidae. They are the only family in the biological order Ciconiiformes, which was once much larger and held a number of families....
s, tropicbird
Tropicbird
Tropicbirds are a family, Phaethontidae, of tropical pelagic seabirds now classified in their own order Phaethontiformes. Their relationship to other living birds is unclear, and they appear to have no close relatives. There are three species in one genus, Phaethon...
s and wader
Wader
Waders, called shorebirds in North America , are members of the order Charadriiformes, excluding the more marine web-footed seabird groups. The latter are the skuas , gulls , terns , skimmers , and auks...
s. If they can be assigned to a modern order (which is highly doubtful), with the possible exception of Marinavis they would probably not be considered Procellariiformes. Eopuffinus and Manu
Manu (genus)
Manu is a genus of prehistoric seabird. It lived during the Early Oligocene, and is known from a few fossil bones found in New Zealand. Its name derives from the Māori language, and is the common Polynesian term for "bird"....
on the other hand are more likely members of the Procellariiformes; the former might be an ancestral petrel, the latter an ancient albatross. As regards Lonchodytes
Lonchodytes
Lonchodytes is a Late Cretaceous genus of aquatic bird, which lived along the shores of the Western Interior Seaway. It lived probably during the Maastrichtian, 70 million years ago , and was found in Lance Creek Formation rocks in Wyoming though it seems still somewhat unclear if it did fossilize...
(or rather its type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
L. estesi), it is the best candidate for the most ancient procellariiform known to date; it pre-dates the evolutionary radiation
Evolutionary radiation
An evolutionary radiation is an increase in taxonomic diversity or morphological disparity, due to adaptive change or the opening of ecospace. Radiations may affect one clade or many, and be rapid or gradual; where they are rapid, and driven by a single lineage's adaptation to their environment,...
that brought about the modern families and hence would occupy a basal position in the order. Parascaniornis on the other hand was formerly assigned to the Procellariiformes by some, but it is actually a hesperornithiform
Hesperornithiformes
Hesperornithes is an extinct and highly specialized clade of Cretaceous toothed birds. Hesperornithine birds, apparently limited to former aquatic habitats in the Northern Hemisphere, include genera such as Hesperornis, Parahesperornis, Baptornis, Enaliornis, and probably Potamornis, all...
synonymous with Baptornis
Baptornis
Baptornis is an extinct genus of flightless aquatic bird from the Late Cretaceous, some 87-80 million years ago . The fossils of Baptornis advenus, the type species, were discovered in Kansas, which at its time was mostly covered by the Western Interior Seaway, a shallow shelf sea...
.
Evolution
Fossil records indicate that Procellariiformes have been around at least 60 million years, but a DNA-based study from 1997 states that they have been around into the Cretaceous Period and survived the Cretaceous–Tertiary extinction event. This Order was distinct from Sphenisciformes, Penguins, and GaviiformesGaviiformes
Gaviiformes is an order of aquatic birds containing the loons or divers and their closest extinct relatives. Modern gaviiformes are found in many parts of North America and northern Eurasia , though prehistoric species were more widespread.-Classification and evolution:There are five living...
, Divers, before the extinction event. Fossil records are rare but 16 million year old fossils show that Albatross
Albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds allied to the procellariids, storm-petrels and diving-petrels in the order Procellariiformes . They range widely in the Southern Ocean and the North Pacific...
es and Shearwater
Shearwater
Shearwaters are medium-sized long-winged seabirds. There are more than 30 species of shearwaters, a few larger ones in the genus Calonectris and many smaller species in the genus Puffinus...
s haven't changed much since then. It is believed that they evolved first in the Southern Hemisphere
Southern Hemisphere
The Southern Hemisphere is the part of Earth that lies south of the equator. The word hemisphere literally means 'half ball' or "half sphere"...
, even though the majority of the fossils have been found in the Northern Hemisphere
Northern Hemisphere
The Northern Hemisphere is the half of a planet that is north of its equator—the word hemisphere literally means “half sphere”. It is also that half of the celestial sphere north of the celestial equator...
. This is likely due to the fact that there is more land to find fossils in the north. DNA evidence has confirmed common ancestry for all Procellariiformes, however, the taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
within the order is complex and fluctuating. The fossil record of the diving petrels goes back to the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
, with a species from that family being described in 2007. The most numerous fossils from the Paleogene are those from the extinct family Diomedeoididae
Diomedeoididae
The Diomedeoididae are a prehistoric family of seabirds. The family was in the order Procellariiformes which today is composed of the albatrosses and petrels. At present the family contains a single genus, Diomedeoides, in which there are three described species...
, fossils of which have been found in Central Europe and Iran.