Procellariformes (Fürbringer, 1888)
Procellariiformes is an order of seabirds that comprises four families: the albatrosses, procellariids, storm-petrels and diving petrels. Formerly called Tubinares and still called tubenoses in English, they are often referred to collectively as the petrels, 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 which 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.
Procellariiformes comes from the Latin 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.
The Procellariiformes have a cosmopolitan distribution across the world's oceans and seas, although at the levels of family and genus there are some clear patterns. Antarctic Petrels, Thalassoica antarctica, have to fly over 100 mi (160 km) to get to the ocean from their breeding colonies in Antarctica, and Northern Fulmars 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 petrels are mostly polar with some temperate species. The majority of the fulmarine petrels, along with the prions, are confined to the southern hemisphere. The shearwaters 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 albatrosses 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 64,000 km (40,000 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 albatrosses 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 Petrels 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 lb) and a 3.6 m (12 ft) wingspan, to the tiny Least Storm-petrel, at 20 g (0.71 oz) and a 32 cm (13 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 anothery unifying feature, although there are differences within the order. Petrels have a plate called Maxillary unguis that forms a hook on their upper bill. The smaller members of the order have a comb-like lower bill, made by the tomia plate, for plankton feeding. Finally, they have a stomach oil stored in their provemtriculus 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 they eyes. This gland 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 in general terms that it removes salt that forms a 5% saline solution that drips out of their nose 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 petrels 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 petrels 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-petrels are cavity nesters, as are many of the procellariids. The fulmarine petrels and some tropical gadfly petrels and shearwaters 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 ft), and a study of Cory's Shearwaters 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 which returned to the same site for 25 years. The average number of birds returning to the same nesting sites is high in all species studied, with figures of around 91% for Bulwer's Petrels, and 85% of males and 76% of females for Cory's Shearwaters (after a successful breeding attempt).
Pair bonds and life history
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 eight or ten years in the case of larger species. Once they begin breeding they make only a single breeding attempt per nesting season, even if the egg is lost early on in the season they will seldom relay. 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.
The majority of Procellariiformes nest once a year and do so seasonally. Some tropical shearwaters, like the Christmas Shearwater, are able to nest on cycles slightly shorter than a year, and the large great albatrosses (genus Diomedea) nest in consecutive years. Most temperate and polar species nest over the spring-summer, although some albatrosses and procellariids nest over the winter. In the tropics some species breed throughout the year, but most nest in discreet periods. Procellariiformes return to the nesting colonies several months before laying, and attend their nesting sites regularly before copulation. Prior to laying females embark on a pre-laying exodus to build up reserves of energy to lay the comparably large egg.
When the female returns and lays the male takes the first incubation stint and the female returns to sea. Incubation is shared between both sexes. The duration of individual stints varies from just a few days to 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 be as long as 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 feathers, 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, for surface nesting species, which have to deal with a greater range of weather and also have to content with predators like skuas and frigatebirds, and consequently have longer guard phases, 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 lipids 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 fledges between two and nine months 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 not a lot of predators at the nesting colonies, therefore there is no pressure to fledge quickly. Second, the time between feedings is long due to the distance and a chick the had a higher growth rate would stand a better chance of starving to death. The durations between feedings vary between species and during the stages of development. Small feeds are frequent during the guard phase, but afterwards become less frequent.
Relationship with humans
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 Coleridge'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 petrels 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.
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 shearwaters and albatrosses along with those of other seabirds in 5,000 year old middens 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 kaitiakitanga. 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.
Threats and conservation
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 Petrels breeding on the Chatham Islands, only 400 Zino's Petrels and only 80 Amsterdam Albatrosses. 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 mammals such as rats, feral cats and pigs. 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 Petrels 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.
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 Keulemans.
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 loons (Gaviiformes) and penguins (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 †Diomedeoididae (Early Oligocene – Early Miocene)
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, cormorants, loons, pelicans, penguins, and perhaps also grebes, flamingos, storks, tropicbirds and waders. 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.
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 Gaviiformes, Divers, before the extinction event. Fossil records are rare but 16 million year old fossils show that Albatrosses and Shearwaters 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.
1. ^ * Brands, Sheila (August 14, 2008). "Systema Naturae 2000 / Classification - Parker (1982)". Project: The Taxonomicon. http://sn2000.taxonomy.nl/Parker/1982/Classification/51469.htm. Retrieved 12 February 2009.
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