- Procellariiformes
Taxobox
name = Procellariiformes
fossil_range = MiddleEocene – Recent
image_width = 250px
image_caption =Cape Petrel "Daption capense"
regnum =Animal ia
phylum = Chordata
classis = Aves
infraclassis =Neognathae
ordo = Procellariiformes
ordo_authority = Fürbringer, 1888
subdivision_ranks = Families
subdivision =Procellariidae Diomedeidae Hydrobatidae
PelecanoididaeProcellariiformes is an order of
seabird s that comprises four families: thealbatross es, procellariids,storm-petrel s anddiving petrel s. Formerly called Tubinares and still called tubenoses in English, they are often referred to collectively as thepetrel s, a term that has been applied to all ProcellariiformesWarham, J. (1996). "The Behaviour, Population, Biology and Physiology of the Petrels". London: Academic Press, ISBN 0-12-735415-8] or more commonly all the families except the albatrosses.Brooke, M. (2004). "Albatrosses And Petrels Across The World" Oxford University Press, Oxford, UK ISBN 0-19-850125-0] They are almost exclusivelypelagic (feeding in the open ocean). They have acosmopolitan distribution across the world's oceans, with the highest diversity being aroundNew 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, marinepollution and the danger of fisheriesby-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 theAgreement on the Conservation of Albatrosses and Petrels , a legally binding international treaty signed in 2001.Biology
Distribution and movements
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. The most cosmopolitan family is theProcellariidae , although within that family there are some gaps in distribution. The gadfly petrels, "Pterodroma ", have a generally tropical and temperate distribution, whereas thefulmarine 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. Theshearwater s have the most widespread distribution, although they are absent from the Pacific north of Japan as breeding birds.The
storm-petrel s are almost as widespread as the procellariids, and fall into two distinct subfamilies; theOceanitinae have a mostly southern hemisphere distribution and theHydrobatinae are found mostly in the northern hemisphere. Amongst thealbatross 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.Olson, S.L., Hearty, P.J. (2003) "Probable extirpation of a breeding colony of Short-tailed Albatross ("Phoebastria albatrus") on Bermuda by Pleistocene sea-level rise." "Proceedings of the National Academy of Science" 100: (22) 12825–12829.] Finally thediving-petrel s 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 offJapan ,Alaska and California, an annual round trip of 64,000 km (40,000 miless), the longest measured annual migration of any bird. [Shaffer S.A., Tremblay Y., Weimerskirch H., Scott D., Thompson D.R., Sagar P.M., Moller H., Taylor G.A., Foley D.G., Block B.A., Costa D.P. (2006) "Migratory shearwaters integrate oceanic resources across the Pacific Ocean in an endless summer." "Proc Natl Acad Sci." 103(34): 12799–12802] A number of otherpetrel species undertake trans-equatorial migrations, including theWilson's Storm-petrel and theProvidence Petrel , but noalbatross es cross do due to their reliance on wind assisted flight. There are other long-distant migrants within the order;Swinhoe's Storm-petrel s breed in the western Pacific and migrates to the western Indian Ocean, [ Van Den Berg AB, Smeenk C, Bosman CAW, Haase BJM, Van Der Niet AM, Cadée GC (1990) "Barau’s petrel "Pterodroma baraui", Jouanin’s petrel "Bulweria fallax" and other seabirds in the northern Indian Ocean in June–July 1984 and 1985." "Ardea" 79:1–14 ] andBonin Petrel s nesting inHawaii migrate to the coast of Japan during the non-breeding season. [Seto, N. W. H., and D. O’Daniel. (1999). Bonin Petrel ("Pterodroma hypoleuca"). In "The Birds of North America", No. 385 (A. Poole and F. Gill, eds.). The Birds of North America, Inc., Philadelphia, PA.]Morphology and flight
Procellariiformes range in size from the very large
Wandering Albatross , at 10 kg, to the tinyLeast Storm-petrel , at 20 g. 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 characteristic tubular nasal passage which is used for
olfaction . [Lequette, B., Verheyden, C., Jowentin, P. (1989) "Olfaction in Subantarctic seabirds: Its phylogenetic and ecological significance" "The Condor" 91: 732-135. [http://scholar.google.com/url?sa=U&q=http://elibrary.unm.edu/sora/Condor/files/issues/v091n03/p0732-p0735.pdf] ] This ability to smell helps to locate patchily distributed prey at sea and may also help locate their nests within nesting colonies. [Bonadonna, Francesco, Cunningham, Gregory B., Jouventin, Pierre, Hesters, Florence, Nevitt, Gabrielle A. (2003)Evidence for nest-odour recognition in two species of diving petrel. J. Exp. Biol. 206:3719-3722 [http://jeb.biologists.org/cgi/content/full/206/20/3719 abstract] ]The longer-winged species fly using a switchback technique to minimise active flapping. All eat fish,
squid or similar marine prey.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.Breeding behaviour
Breeding colonies
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 ofLeach's Storm Petrel s. [West, J. & Nilsson, R. (1994) "Habitat use and burrow densities of burrow-nesting seabirds on South East Island, Chatham Islands, New Zealand" "Notornis (Supplement)" 41 27-37 [http://www.notornis.org.nz/free_issues/Notornis_41-1994/Notornis_41_sup_27.pdf#search=%22storm-petrel%22] ] For almost all species the need to breed is the only reason that Procellariiformes return to land at all. 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.Medeiros R., Hothersall B. and Campos A. (2003) "The use of artificial breeding chambers as a conservation measure for cavity-nesting procellariiform seabirds: a case study of the Madeiran storm petrel ("Oceanodroma castro")" "Biological Conservation " 116(1): 73-80 http://dx.doi.org/10.1016/S0006-3207(03)00178-2] Burrows and natural crevices are most commonly used by the smaller species; all thestorm-petrel s anddiving-petrel s are cavity nesters, as are many of theprocellariid s. Thefulmarine petrel s and some tropicalgadfly petrel s andshearwater 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 andmitochondrial 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; inLaysan Albatross the average distance between hatching site and the site where a bird established its own territory was 22 metres, [Fisher, H.I., (1976) "Some dynamics of a breeding colony of Laysan Albatrosses. "Wilson Bulletin" 88: 121–142.] and a study ofCory's Shearwater s nesting nearCorsica 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. [Rabouam, C., Thibault, J.-C., Bretagnolle, V., (1998) "Natal Philopatry and Close Inbreeding in Cory's Shearwater ("Calonectris diomedea")" "Auk" 115(2): 483–486 [http://elibrary.unm.edu/sora/Auk/v115n02/p0483-p0486.pdf] ]Mitochondrial DNA provides evidence of restrictedgene flow between different colonies, strongly suggesting philopatry. [Ovenden, J.R., Wust-Saucy, A., Bywater, R., Brothers, N., White, R.W.G. (1991) " Genetic evidence for philopatry in a colonially nesting seabird. the Fairy Prion ("Pachyptila turtur")" "Auk" 108: 688–694. [http://scholar.google.com/url?sa=U&q=http://elibrary.unm.edu/sora/Auk/v108n03/p0688-p0694.pdf] ]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% forBulwer's Petrel s, [Mouguin, J-L., (1996) "Faithfulness to mate and nest site of Bulwer's Petrel, "Bulweria bulweria" at Selvagem Grande" "Marine Ornithology" 24: 15–18. [http://www.marineornithology.org/PDF/24/24_2.pdf] ] and 85% of males and 76% of females forCory's Shearwater s (after a successful breeding attempt). [ Thibault, J-C., (1994) "Nest-site tenacity and mate fidelity in relation to breeding success in Cory's Shearwater "Calonectris diomedea" "Bird Study" 41(1): 25–28.]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. [Jouventin, P., Monicault, G. de & Blosseville, J.M. (1981) "La danse de l'albatros, "Phoebetria fusca". "Behaviour" 78: 43–80.] 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). [Pickering, S.P.C., & Berrow, S.D., (2001) "Courtship behaviour of the Wandering Albatross "Diomedea exulans" at Bird Island, South Georgia" "Marine Ornithology" 29: 29–37 [http://www.marineornithology.org/PDF/29_1/29_1_6.pdf] ] 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. [ Genevois, F; Bretagnolle, V (1994) "Male Blue Petrels reveal their body mass when calling" "Ethology Ecology & Evolution" 6(3): 377–383.] 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, [Robertson, C.J.R. (1993). "Survival and longevity of the Northern Royal Albatross "Diomedea epomophora sanfordi" at Taiaroa Head" 1937–93. "Emu" 93: 269–276.] even the tiny storm-petrels are known to have survived for 30 years. [Klimkiewicz, M. K. 2007. [http://www.pwrc.usgs.gov/BBL/homepage/longvrec.htm Longevity Records of North American Birds] . Version 2007.1. Patuxent Wildlife Research Center. Bird Banding Laboratory. Laurel MD.]
Nesting and chick rearing
The majority of Procellariiformes nest once a year and do so seasonally. [Brooke, M. (2004) p.46 ] Some tropical shearwaters, like the
Christmas Shearwater , are able to nest on cycles slightly shorter than a year, and the largegreat albatross es (genus "Diomedea ") nest in consecutive years. Most temperate and polar species nest over the spring-summer , although some albatrosses and procellariids nest over thewinter . 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. [Warham, J. (1990) "The Petrels - Their Ecology and Breeding Systems" London: Academic Press. ISBN 978-0127354200] 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. [Brooke, M. (2004) p.67 ]
Upon hatching the chicks are semi-
precocial , having open eyes, a dense covering of white or greydown 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, for surface nesting species, which have to deal with a greater range of weather and also have to content with predators likeskua s andfrigatebird s, and consequently have longer guard phases, as long as two weeks in procellariids and three weeks in albatrosses. [Brooke, M. (2004) p.75 ] The chick is fed by both parents. Chicks are fed on fish, squid, krill andstomach oil . Stomach oil isoil composed of neutral dietarylipid s that are the residue created bydigestion 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 fordiesel oil. [Warham, J. (1976) "The Incidence, Function and ecological significance of petrel stomach oils." "Proceedings of the New Zealand Ecological Society" 24 84-93 [http://www.nzes.org.nz/nzje/free_issues/ProNZES24_84.pdf] ] This can be a real advantage for species that range over huge distances to provide food for hungry chicks. [Roby, Daniel D, Taylor, Jan R E, Place, Allen R (1997) "Significance of stomach oil for reproduction in seabirds: An interspecies cross-fostering experiment." "The Auk" 114 (4) 725-736. [http://www.findarticles.com/p/articles/mi_qa3793/is_199710/ai_n8765697] ] The oil is also used in defence. All Procellariiformes create stomach oil except thediving-petrel s.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
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".Carboneras, C. (1992) "Family Diomedeidae (Albatross)" in "Handbook of Birds of the World" Vol 1. Barcelona:Lynx Edicions, ISBN 84-87334-10-5] 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. [Lasky E (1992). "A Modern Day Albatross: The Valdez and Some of Life's Other Spills." "The English Journal", 81 (3): 44–46. DOI|10.2307/820195] There are few instances ofpetrel s in culture, although there are sailors legends regarding thestorm-petrel s, 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.Carboneras, C. (1992) "Family Hydrobatidae (Storm-petrels)" P.p 258-265 in "Handbook of Birds of the World" Vol 1. Barcelona:Lynx Edicions, ISBN 84-87334-10-5]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
shearwater s and albatrosses along with those of other seabirds in 5,000 year oldmidden s inChile , [Simeone, A. & Navarro, X. (2002) "Human exploitation of seabirds in coastal southern Chile during the mid-Holocene" "Rev. chil. hist. nat" 75(2): 423–431 [http://www.scielo.cl/scielo.php?script=sci_arttext&pid=S0716-078X2002000200012&lng=es&nrm=iso&tlng=en] ] 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 toextinction . Some cultures continue to harvest shearwaters (a practice known asmuttonbirding ); for example theMaori ofNew Zealand ,who use a sustainable traditional method known as "kaitiaki tanga".Threats and conservation
Exotic species introduced to the remote breeding colonies is also a threat to all types of Procellariiformes. These principally take the form of
predator s; most albatross and petrel species are clumsy on land and are unable to defend themselves frommammal s such asrat s,feral cat s andpig s. This phenomenon, known as ecological naivete, has resulted in numerous declines in many species and has been strongly implicated in theextinction of theGuadalupe Storm-petrel . [A contemporary account of the decline of the Guadalupe Storm-petrel can be found here - Thayer, J. & Bangs, O (1908) "The Present State of the Ornis of Guadaloupe Island" "Condor" 10(3): 101-106 [http://elibrary.unm.edu/sora/Condor/files/issues/v010n03/p0101-p0106.pdf#search=%22The%20Present%20State%20of%20the%20Ornis%20of%20Guadaloupe%20Island%20%22] ] Introduced herbivores can also cause problems if they unbalance theecology of the island; introduced rabbits destroyed the forest understory on Cabbage Tree Island offNew South Wales ; this both increased the vulnerability of theGould'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.Carlile, N., Proiddel, D., Zino, F., Natividad, C. & Wingate, D.B. (2003) "A review of four successful recovery programmes for threatened sub-tropical petrels" "Marine Ornithology" 31: 185–192] In the past exploitation was a threat (see above), although this is less of a threat now. Other threats the ingestion of plasticflotsam . 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. [Pierce, K., Harris, R., Larned, L., Pokras, M., (2004) "Obstruction and starvation associated with plastic ingestion in a Northern Gannet "Morus bassanus" and a Greater Shearwater "Puffinus gravis" "Marine Ornithology" 32: 187–189 [http://www.marineornithology.org/PDF/32_2/32_2_187-189.pdf] ] This can also be picked up by foraging adults and fed to chicks, stunting their development and reducing the chances of successfully fledging. [Auman, H.J., Ludwig, J.P., Giesy, J.P., Colborn, T., (1997) "Plastic ingestion by Laysan Albatross chicks on Sand Island, Midway Atoll, in 1994 and 1995." in "Albatross Biology and Conservation", (ed by G. Robinson and R. Gales). Surrey Beatty & Sons: Chipping Norton. Pp. 239-44 [http://www.mindfully.org/Plastic/Ocean/Albatross-Plastic-Ingestion1997.htm] ] Procellariids are also vulnerable to general marine pollution, as well as oil spills. Some species, such as theBarau's Petrel and theNewell's Shearwater , which nest high up on large developed islands are victims of light pollution. Chicks that are fledging are attracted to streetlights and are unable to reach the sea. An estimated 20–40% of fledging Barau's Petrels are attracted to the streetlights onRéunion . [Le Correa, M., Ollivier, A., Ribesc S., Jouventin, P., (2002) " [http://web.archive.org/web/20061116215442/http://www.cefe.cnrs.fr/ecomp/PDF+papers/LeCorre+2002.pdf Light-induced mortality of petrels: a 4-year study from Réunion Island (Indian Ocean)] " "Biological Conservation" 105: 93–102]pecies
There are a total of around 125 species of Procellariiformes world-wide, divided among four families:
* Order Procellariiformes
** FamilyProcellariidae (shearwater s,fulmarine petrel s,gadfly petrel s, and prions)
** FamilyDiomedeidae (albatross es)
** FamilyHydrobatidae (storm-petrel s)
** FamilyPelecanoididae (diving-petrel s)The Hydrobatidae's two subfamilies, Oceanitinadae and Hydrobatinae, are probably better treated as distinct families. [Nunn, G & Stanley, S. (1998): Body Size Effects and Rates of Cytochrome "b" Evolution in Tube-Nosed Seabirds. "Molecular Biology and Evolution" 15(10): 1360-1371 [http://mbe.oxfordjournals.org/cgi/reprint/15/10/1360.pdf PDF fulltext] [http://mbe.oxfordjournals.org/cgi/reprint/17/11/1774.pdf Corrigendum] ]
"
Primodroma ", a fossil from theEarly Eocene London Clay ofEngland , may belong to the Hydrobatidae.Procellariiformes are very closely related to
Sphenisciformes (Penguins).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 may be correctFact|date=February 2007.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 .References
*
External links
* [http://www.acap.aq/ The Agreement for the Conservation of Albatrosses and Petrels (ACAP)]
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