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Procellariidae
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{{Short description|Family of seabirds which includes petrels, shearweters and prions }} {{pp-move|small=yes}} {{Automatic taxobox | name = Procellariidae | image = Damier du Cap - Cape Petrel.jpg | image_caption = [[Cape petrel]] (''Daption capense'') | taxon = Procellariidae | diversity = 16 genera, 99 species, of which 3 are extinct | authority = [[William Elford Leach|Leach]], 1820 | subdivision_ranks = Genera | subdivision = ''[[Macronectes]]''<br /> ''[[Fulmarus]]''<br /> ''[[Thalassoica]]''<br /> ''[[Daption]]''<br /> ''[[Snow petrel|Pagodroma]]''<br /> ''[[Halobaena]]''<br /> ''[[Pachyptila]]''<br /> ''[[Procellaria]]''<br /> ''[[Bulweria]]''<br /> ''[[Calonectris]]''<br /> ''[[Puffinus]]''<br /> ''[[Pelecanoides]]''<br /> ''[[Ardenna]]''<br /> ''[[Pseudobulweria]]''<br /> ''[[Aphrodroma]]''<br /> ''[[Pterodroma]]'' }} The [[family (biology)|family]] '''Procellariidae''' is a group of [[seabird]]s that comprises the [[fulmarine petrel]]s, the [[gadfly petrel]]s, the [[diving petrel]]s, the [[prion (bird)|prions]], and the [[shearwater]]s. This family is part of the bird [[order (biology)|order]] [[Procellariiformes]] (or tubenoses), which also includes the [[albatross]]es and the [[storm petrel]]s. The procellariids are the most numerous family of tubenoses, and the most diverse. They range in size from the [[giant petrel]]s with a wingspan of around {{cvt|2.0|m}}, that are almost as large as the albatrosses, to the diving petrels with a wingspan of around {{cvt|34|cm}} that are similar in size to the [[little auk]]s or dovekies in the family [[Alcidae]]. Male and female birds are identical in appearance. The plumage color is generally dull, with blacks, whites, browns and grays. The birds feed on fish, [[squid]] and [[crustacea]], with many also taking [[Discards|fisheries discards]] and [[carrion]]. Whilst agile swimmers and excellent in water, petrels have weak legs and can only shuffle on land, with the giant petrels of the genus ''[[Macronectes]]'' being the only two species that are capable of proper [[terrestrial locomotion]]. All species are accomplished long-distance foragers, and many undertake long [[equator|trans-equatorial]] [[bird migration|migrations]]. They are [[seabird colony|colonial breeders]], exhibiting long-term [[Monogamy in animals|mate fidelity]] and [[philopatry|site philopatry]]. In all species, a single white egg is laid each breeding season. The parents take it in turns to incubate the egg and to forage for food. The feeding area can be at a great distance from the nest site. The incubation times and chick-rearing periods are exceptionally long compared to other birds. Many procellariids have breeding populations of over several million pairs; others number fewer than 200 birds. Humans have traditionally exploited several species of [[fulmar]] and shearwater (known as muttonbirds) for food, fuel, and bait, a practice that continues in a controlled fashion today. Several species are threatened by [[introduced species]] attacking adults and chicks in breeding colonies and by [[long-line fishing|long-line fisheries]]. ==Taxonomy and evolution== The [[family (biology)|family]] Procellariidae was introduced (as Procellaridæ) by the English zoologist [[William Elford Leach]] in a guide to the contents of the [[British Museum]] published in 1820.<ref>{{ cite book | last=Leach | first=William Elford | author-link=William Elford Leach | year=1820 | chapter=Eleventh Room | title=Synopsis of the Contents of the British Museum | volume=17 | place=London | publisher=British Museum | edition=17th| page=68 | url=https://www.biodiversitylibrary.org/page/55257931 }} Although the name of the author is not specified in the document, Leach was the Keeper of Zoology at the time.</ref><ref>{{cite book | last=Bock | first=Walter J. | year=1994 | title=History and Nomenclature of Avian Family-Group Names | series=Bulletin of the American Museum of Natural History | volume= 222 | publisher=American Museum of Natural History | place=New York | pages=110, 130, 161 | hdl=2246/830 }}<!--Linked page allows download of the 48MB pdf--></ref> The name is derived from the [[type genus]] ''[[Procellaria]]'' which in turn is derived from the [[Latin]] word ''procella'' meaning "storm" or "gale".<ref>{{cite book | last=Jobling | first=James A. | year=2010| title=The Helm Dictionary of Scientific Bird Names | publisher=Christopher Helm | location=London | isbn=978-1-4081-2501-4 | page=317 | url=https://archive.org/stream/Helm_Dictionary_of_Scientific_Bird_Names_by_James_A._Jobling#page/n317/mode/1up }}</ref> The type genus was named in 1758 by the Swedish naturalist [[Carl Linnaeus]] in the [[10th edition of Systema Naturae|tenth edition]] of his ''[[Systema Naturae]]''.<ref>{{cite book | last=Linnaeus | first=Carl | authorlink=Carl Linnaeus | year=1758 | title= Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis | volume=1 | edition=10th | page=131 | publisher=Laurentii Salvii | place=Holmiae (Stockholm) | language=Latin | url=https://www.biodiversitylibrary.org/page/727036 }}</ref> Procellariidae is one of families that make up the [[Order (biology)|order]] [[Procellariiformes]].<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | date=July 2021 | title=Petrels, albatrosses | work=IOC World Bird List Version 11.2 | url=https://www.worldbirdnames.org/bow/petrels/ | publisher=International Ornithologists' Union | access-date=15 January 2022 }}</ref> Before the introduction of [[molecular phylogenetic]]s, the traditional arrangement was to divide the Procellariiformes into a set of four families: [[Diomedeidae]] containing the albatrosses, [[Hydrobatidae]] containing all the storm petrels, Pelecanoididae containing the [[diving petrel]]s and Procellariidae containing the petrels, shearwaters and fulmars.<ref>{{ cite book | editor1-last=Mayr | editor1-first=Ernst | editor1-link=Ernst Mayr | editor2-last=Cottrell | editor2-first=G. William | year=1979 | title=Check-List of Birds of the World | volume=1 | edition=2nd | publisher=Museum of Comparative Zoology | location=Cambridge, Massachusetts | pages=48–121 | url=https://www.biodiversitylibrary.org/item/54612 }}</ref><ref name=hanzaborder>{{cite book | year=1990 | chapter=Order Procellariiformes | editor1-last=Marchant | editor1-first=S. | editor2-last=Higgins | editor2-first=P.G. | title=Handbook of Australian, New Zealand & Antarctic Birds. Volume 1: Ratites to ducks; Part A, Ratites to petrels | location=Melbourne, Victoria | publisher=Oxford University Press | isbn=978-0-19-553068-1 | pages=263–264 }}</ref> The family Hydrobatidae was further divided into two [[subfamilies]], the northern storm petrels in Hydrobatinae and the southern or austral storm petrels in Oceanitinae. A 1998 analysis of mitochondrial [[cytochrome b|cytochrome ''b'']] sequences found there was deep genetic divergence between the two subfamilies.<ref name="Nunn"/> Subsequent large-scale multigene studies found that the two subfamilies were not [[sister taxon|sister taxa]].<ref>{{ cite journal | last1=Hackett | first1=S.J.| last2=Kimball | first2=R.T. | last3=Reddy | first3=S. | last4=Bowie | first4=R.C.K.| last5=Braun | first5=E.L.| last6=Braun | first6=M.J.| last7=Chojnowski | first7=J.L.| last8=Cox | first8=W.A. | last9=Han | first9=K-L. | last10=Harshman | first10=J. | last11=Huddleston | first11=C.J.| last12=Marks | first12=B.D.| last13=Miglia | first13=K.J.| last14=Moore | first14=W.S.| last15=Sheldon | first15=F.H.| last16=Steadman | first16=D.W.| last17=Witt | first17=C.C.| last18= Yuri | first18= T. | year=2008 | title=A phylogenomic study of birds reveals their evolutionary history | journal=Science | volume=320 | issue=5884 | pages=1763–1767 | doi=10.1126/science.1157704 | pmid=18583609| bibcode=2008Sci...320.1763H| s2cid=6472805}}</ref><ref name=prum>{{ cite journal | last1=Prum | first1=R.O. | author1-link=Richard Prum | last2=Berv | first2=J.S. | last3=Dornburg | first3=A. | last4=Field | first4=D.J. | last5=Townsend | first5=J.P. | last6=Lemmon | first6=E.M. | last7=Lemmon | first7=A.R. | year=2015 | title=A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing | journal=Nature | volume=526 | issue=7574 | pages=569–573 | doi=10.1038/nature15697 | pmid=26444237 | bibcode=2015Natur.526..569P | doi-access= | s2cid=205246158 }}</ref><ref name=kuhl>{{Cite journal | last1=Kuhl | first1=H. | last2=Frankl-Vilches | first2=C. | last3=Bakker | first3=A. | last4=Mayr | first4=G. | last5=Nikolaus | first5=G. | last6=Boerno | first6=S.T. | last7=Klages | first7=S. | last8=Timmermann | first8=B. | last9=Gahr | first9=M. | date=2020 | title=An unbiased molecular approach using 3′-UTRs resolves the avian family-level tree of life | journal=Molecular Biology and Evolution | volume=38 | issue=msaa191 | pages=108–127 | doi=10.1093/molbev/msaa191 | pmid=32781465 | pmc=7783168 | doi-access=free }}</ref> The storm petrels were therefore split into two families: Hydrobatidae containing the northern storm petrels and Oceanitidae, containing the southern storm petrels.<ref name=ioc/> The multigene genetic studies found that the diving petrels in the family Pelecanoididae were nested within the family Procellariidae.<ref name=prum/><ref name=kuhl/><ref name=reddy>{{Cite journal | last1=Reddy | first1=S. | last2=Kimball | first2=R.T. | last3=Pandey | first3=A. | last4=Hosner | first4=P.A. | last5=Braun | first5=M.J. | last6=Hackett | first6=S.J. | last7=Han | first7=K.-L. | last8=Harshman | first8=J. | last9=Huddleston | first9=C.J. | last10=Kingston | first10=S. | last11=Marks | first11=B.D. | last12=Miglia | first12=K.J. | last13=Moore | first13=W.S. | last14=Sheldon | first14=F.H. | last15=Witt | first15=C.C. | last16=Yuri | first16=T. | last17=Braun | first17=E.L. | date=2017 | title=Why do phylogenomic data sets yield conflicting trees? Data type influences the avian tree of life more than taxon sampling | journal=Systematic Biology | volume=66 | issue=5 | pages=857–879 | doi=10.1093/sysbio/syx041 | pmid=28369655 | doi-access=free }}</ref> As a result, the diving petrels was merged into Procellariidae.<ref name=ioc/> The molecular evidence suggests that the albatrosses were the first to diverge from the ancestral stock, and the [[Oceanitidae|austral storm petrels]] next, with the procellariids and northern storm petrels splitting most recently.<ref name=prum/><ref name=kuhl/><ref name=reddy/> {{Clade|label1=Procellariiformes |1={{clade |1=[[Diomedeidae]] – albatrosses (21 species) |2={{clade |1=[[Oceanitidae]] – austral storm petrels (9 species) |2={{clade |1=[[Hydrobatidae]] – northern storm petrels (18 species) |2='''Procellariidae''' – petrels and shearwaters (99 species) }} }} }} }} Within the procellariid family, a genetic analysis based on the cytochrome ''b'' gene published in 2004 indicated that the genus ''Puffinus'' contained two distinct [[clades]] and was [[polyphyletic]].<ref name=Penhallurick>{{cite journal | last1=Penhallurick | first1=John | last2=Wink | first2=Michael | year=2004 | title=Analysis of the taxonomy and nomenclature of the Procellariiformes based on complete nucleotide sequences of the mitochondrial cytochrome ''b'' gene | journal=Emu |volume= 104 | issue=2 | pages= 125–147 | doi=10.1071/MU01060| bibcode=2004EmuAO.104..125P | s2cid=83202756 }}</ref> The genus was therefore split and a group of species moved into the resurrected genus ''[[Ardenna]]''.<ref name=ioc/><ref>{{ cite web | last=Remsen | first=J.V. | title=Proposal (647) to South American Classification Committee: Split ''Ardenna'' from ''Puffinus'' | publisher=South American Classification Committee | date=September 2014 | url=http://www.museum.lsu.edu/~Remsen/SACCprop647.htm | accessdate=23 January 2016}}</ref> The other genera within the family were found to be [[monotypic]] but the relationships between the genera remained unclear.<ref name=Penhallurick/> This changed when a multigene genetic study published in 2021 provided a genus-level phylogeny of the family.<ref name=estandia>{{Cite journal | last1=Estandía | first1=A. | last2=Chesser | first2=R.T. | last3=James | first3=H.F. | last4=Levy | first4=M.A. | last5=Ferrer Obiol | first5=J. | last6=Bretagnolle | first6=V. | last7=González-Solís | first7=J. | last8=Welch | first8=A.J. | accessdate=11 December 2021 | date=27 July 2021 | title=Substitution rate variation in a robust Procellariiform seabird phylogeny is not solely explained by body mass, flight efficiency, population size or life history traits | journal=bioRxiv | url=https://www.biorxiv.org/content/10.1101/2021.07.27.453752v1 | pages=2021.07.27.453752| doi=10.1101/2021.07.27.453752 | s2cid=236502443 }}</ref> {{Clade |label1='''Procellariidae''' |1={{clade |1={{clade |1=''[[Snow petrel|Pagodroma]]'' – snow petrel |2={{clade |1=''[[Thalassoica]]'' – Antarctic petrel |2={{clade |1=''[[Daption]]'' – Cape petrel |2={{clade |1=''[[Macronectes]]'' – giant petrels (2 species) |2=''[[Fulmarus]]'' – fulmars (2 species) }} }} }} }} |2={{clade |1={{clade |1={{clade |1=''[[Pelecanoides]]'' – diving petrels (4 species) |2={{clade |1=''[[Halobaena]]'' – blue petrel |2=''[[Pachyptila]]'' – prions (7 species) }} }} |2={{clade |1=''[[Aphrodroma]]'' – Kerguelen petrel |2=''[[Pterodroma]]'' – gadfly petrels (35 species) }} }} |2={{clade |1={{clade |1=''[[Bulweria]]'' – petrels (2 extant species) |2=''[[Pseudobulweria]]'' – petrels (4 species) }} |2={{clade |1=''[[Procellaria]]'' – petrels (5 species) |2={{clade |1=''[[Puffinus]]'' – shearwaters (21 species) |2={{clade |1=''[[Calonectris]]'' – shearwaters (4 species) |2=''[[Ardenna]]'' – shearwaters (7 species) }} }} }} }} }} }} }} There are 99 [[species]] of procellariid in 16 [[genus|genera]].<ref name=ioc/> The family has usually been broken up into four fairly distinct groups; the [[fulmarine petrel]]s, the [[gadfly petrel]]s, the [[prion (bird)|prions]], and the [[shearwater]]s. With the inclusion of the diving petrels there are now five main groups.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/216/mode/1up 216]}}<ref name=hanzabfam>{{cite book | year=1990 | chapter=Family Procellariidae: fulmars, petrels, prions, shearwaters | editor1-last=Marchant | editor1-first=S. | editor2-last=Higgins | editor2-first=P.G. | title=Handbook of Australian, New Zealand & Antarctic Birds. Volume 1: Ratites to ducks; Part A, Ratites to petrels | location=Melbourne, Victoria | publisher=Oxford University Press | isbn=978-0-19-553068-1 | pages=355–356 }}</ref> * The fulmarine petrels include the largest procellariids, the [[giant petrel]]s, as well as the two [[fulmar]] species, the [[snow petrel]], the [[Antarctic petrel]], and the [[Cape petrel]]. The fulmarine petrels are a diverse group with differing habits and appearances, but are linked [[morphology (biology)|morphologically]] by their [[skull]] features, particularly the long prominent nasal tubes.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/218/mode/1up 218]}} *The four diving petrels are the smallest procellariids with lengths of around {{cvt|20|cm}} and wingspans of {{cvt|33|cm}}. They are compact birds with short wings that are adapted for use under water. They have a characteristic whirring flight and dive into the water without settling. They probably remain all year in the seas near their breeding sites.<ref name=hanzabpele>{{cite book | year=1990 | chapter=Family Pelecanoididae: diving-petrels | editor1-last=Marchant | editor1-first=S. | editor2-last=Higgins | editor2-first=P.G. | title=Handbook of Australian, New Zealand & Antarctic Birds. Volume 1: Ratites to ducks; Part A, Ratites to petrels | location=Melbourne, Victoria | publisher=Oxford University Press | isbn=978-0-19-553068-1 | page=719 }}</ref> * The gadfly petrels, so named due to their helter-skelter flight, are the 35 species in the [[genus]] ''[[Pterodroma]]''. The species vary from small to medium sizes, {{cvt|26|–|46|cm}} in length, and are long winged with short hooked bills.<ref name=Imber1985>{{cite journal | last=Imber | first=M.J. | year = 1985 | title = Origins, phylogeny and taxonomy of the gadfly petrels ''Pterodroma spp'' | journal = Ibis | volume = 127 | issue = 2| pages = 197–229 | doi=10.1111/j.1474-919x.1985.tb05055.x}}</ref> They are most closely related to [[Kerguelen petrel]] which is placed in its own genus ''Aphrodroma''.<ref name=estandia/> * The prions comprise seven species of true prion in the genus ''[[Pachyptila]]'' and the closely related [[blue petrel]]. Often known in the past as whalebirds, three species have large bills filled with [[Lamellae (zoology)|lamellae]] that they use to filter [[plankton]] somewhat as [[baleen whale]]s do, though the old name derives from their association with whales, not their bills (though "prions" does, deriving from [[Ancient Greek]] for "[[saw]]"). They are small procellariids, {{cvt|25|–|30|cm}} in length, with a prominent dark M-shaped mark across the upperwing of their grey plumage. All are restricted to the southern hemisphere.{{sfn|Carboneras|1992a|pp=[https://archive.org/details/handbookofbirdso0001unse/page/219/mode/1up 219], [https://archive.org/details/handbookofbirdso0001unse/page/223/mode/1up 223], [https://archive.org/details/handbookofbirdso0001unse/page/248/mode/1up 248–250]}} * The shearwaters are adapted for diving after prey instead of foraging on the ocean's surface; several species have been recorded diving deeper than {{cvt|30|m}}.<ref>{{Cite journal | last1=Shoji | first1=A. | last2=Dean | first2=B. | last3=Kirk | first3=H. | last4=Freeman | first4=R. | last5=Perrins | first5=C.M. | last6=Guilford | first6=T. | date=2016 | title=The diving behaviour of the Manx Shearwater ''Puffinus puffinus'' | journal=Ibis | volume=158 | issue=3 | pages=598–606 | doi=10.1111/ibi.12381| url=https://discovery.ucl.ac.uk/id/eprint/10075136/ | url-access=subscription }}</ref> They are known for the long trans-equatorial migrations undertaken by many species.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/221/mode/1up 221]}} The shearwaters include the 20 or so species of the genus ''[[Puffinus]]'', seven species in the genus ''[[Ardenna]]'', as well as the five large ''[[Procellaria]]'' species and the four ''[[Calonectris]]'' species. While all these four genera are often known collectively as shearwaters, the ''Procellaria'' are called petrels in their common names.<ref name=ioc/> ==Morphology and flight== [[Image:Giant petrel - Macronectes giganteus.jpg|thumb|left|The flight of [[giant petrel]]s is aided by a shoulder-lock that holds their wing out without effort.| alt=Photo of a giant petrel in flight]] [[Image:Christmas shearwater takeoff.JPG|thumb|alt=Dark brown bird with outstretched wings prepares to take off from sandy beach|left|To take off this [[Christmas shearwater]] (''Puffinus nativitatis'') must face into a strong wind. In calm conditions it must run in order to obtain a high airspeed.]] The procellariids are small- to medium-sized seabirds. The largest, the [[southern giant petrel]] with a wingspan of {{cvt|185|to|205|cm}}, is almost as large as albatrosses; the smallest, the diving petrels have a wingspan of {{cvt|30|to|38|cm}} and are similar in size to [[little auk]]s or dovekies in the family [[Alcidae]].{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/235/mode/1up 235]}}{{sfn|Carboneras|1992b|pp=[https://archive.org/details/handbookofbirdso0001unse/page/272/mode/1up 277–278]}}<ref>{{ cite book | last=Nettleship | first=D.N. | year=1996 | chapter=Family Alcidae (Auks) | editor1-last=del Hoyo | editor1-first=J. | editor2-last=Elliott | editor2-first=A. | editor3-last=Sargatal | editor3-first=J. | title=Handbook of the Birds of the World | volume=3: Hoatzin to Auks | location=Barcelona, Spain | publisher=Lynx Edicions | isbn=978-84-87334-20-7 | pages=678–722 [709] | chapter-url=https://archive.org/details/handbookofbirdso0003unse/page/709/mode/1up | chapter-url-access=registration }}</ref> There are no obvious differences between the sexes, although females tend to be slighter.{{sfn|Brooke|2004|p=5}}{{sfn|Warham|1990|p=4}}<!--haven't found male larger in Brooke--> Like all Procellariiformes, the procellariids have a characteristic tubular nasal passage used for olfaction.<ref name=j2/> This ability to smell helps to locate patchily distributed prey at sea and may help locate [[seabird colony|nesting colonies]]. The [[plumage]] of the procellariids is usually dull, with greys, bluish greys, blacks and browns being the usual colours,{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/218/mode/1up 218]}} although some species have striking patterns such as the [[Cape petrel]].{{sfn|Carboneras|1992a|pp=[https://archive.org/details/handbookofbirdso0001unse/page/240/mode/1up 240–241]}} The technique of [[bird flight|flight]] among procellariids depends on foraging methods. Compared to an average bird, all procellariids have a high [[aspect ratio]] (meaning their wings are long and narrow) and a heavy [[wing loading]]. Therefore, they must maintain a high speed in order to remain in the air. Most procellariids use two techniques to do this, namely, [[dynamic soaring]] and [[slope soaring]].<ref name ="pen"/> Dynamic soaring involves gliding across wave fronts, thus taking advantage of the vertical [[wind gradient]] and minimising the effort required to stay in the air. Slope soaring is more straightforward: the procellariid turns to the wind, gaining height, from where it can then glide back down to the sea. Most procellariids aid their flight by means of flap-glides, where bursts of flapping are followed by a period of gliding; the amount of flapping dependent on the strength of the wind and the choppiness of the water.{{sfn|Warham|1996|p=406}} <!--Shearwaters and other larger petrels, which have lower aspect ratio, must make more use of flapping to remain airborne than gadfly petrels.--> Because of the high speeds required for flight, procellariids need to either run or face into a strong wind in order to take off.{{sfn|Brooke|2004|p=29}} The giant petrels share with the albatrosses an adaptation known as a shoulder-lock: a sheet of [[tendon]] that locks the wing when fully extended, allowing the wing to be kept up and out without any muscle effort.<ref name="pen"/> Gadfly petrels often feed on the wing, snapping prey without landing on the water. The flight of the smaller prions is similar to that of the [[storm petrel]]s, being highly erratic and involving weaving and even looping the loop. The wings of all species are long and stiff. In some species of shearwater the wings are used to power the birds underwater while diving for prey. Their heavier wing loadings, in comparison with surface-feeding procellariids, allow these shearwaters to achieve considerable depths (below {{convert|70|m|ft|abbr=on}} in the case of the [[short-tailed shearwater]]).<ref name=j19/> Procellariids generally have weak legs that are set back, and many species move around on land by resting on the breast and pushing themselves forward, often with the help of their wings.{{sfn|Warham|1996|pp=390–391}} The exceptions to this are the two species of giant petrel, which have strong legs used when they feed on land.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/218/mode/1up 218]}} ==Distribution and migration== {{See also|List of Procellariiformes by population}} [[Image:Shearwater migrant flock.jpg|thumb|Million-strong flocks of shearwaters migrate from New Zealand to Alaska every year. | alt=Photo of a flock of shearwaters in flight]] The procellariids are present in all the world's oceans and most of the seas. They are absent from the [[Bay of Bengal]] and [[Hudson Bay]], but are present year round or seasonally in the rest. The seas north of [[New Zealand]] are the centre of procellariid [[biodiversity]], with the most species.<ref name = "Burger">{{cite book | editor1-last=Schreiber | editor1-first=Elizabeth A. | editor2-last=Burger | editor2-first=Joanne | year=2002 | title=Biology of Marine Birds | location=Boca Raton | publisher=CRC Press | isbn=978-1-4200-3630-5 | page=63 }}</ref><ref>{{ cite journal | last1=Chown | first1=S.L. | last2=Gaston | first2=K.J. | last3=Williams | first3=P.H. | year=1998 | title=Global patterns in species richness of pelagic seabirds: the Procellariiformes | journal=Ecography | volume=21 | issue=4 | pages=342–350 | doi=10.1111/j.1600-0587.1998.tb00399.x | doi-access= | bibcode=1998Ecogr..21..342C }}</ref> Among the groups, the fulmarine petrels have a mostly [[Polar region|polar]] distribution, with most species living around [[Antarctica]] and one, the [[fulmar|northern fulmar]] ranging in the Northern [[Atlantic]] and [[Pacific]] Oceans.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/221/mode/1up 221]}} Of the four species of diving petrel, two are found along the coasts of South America, while the remaining two have circumpolar distributions in the Southern Ocean.{{sfn|Carboneras|1992b|p=[https://archive.org/details/handbookofbirdso0001unse/page/272/mode/1up 272]}} The prions are restricted to the [[Southern Ocean]], and the gadfly petrels are found mostly in the tropics with some temperate species. The shearwaters are the most widespread group and breed in most temperate and tropical seas.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/221/mode/1up 221]}} Many procellariids undertake long annual [[bird migration|migrations]] in the non-breeding season. Southern species of shearwater such as the [[sooty shearwater]] and short-tailed shearwater, breeding on islands off [[Australia]], New Zealand and [[Chile]], undertake transequatorial migrations of millions of birds up to the waters off [[Alaska]] and back each year during the austral winter.<ref>{{Cite journal | last1=Shaffer | first1=S.A. | last2=Tremblay | first2=Y. | last3=Weimerskirch | first3=H. | last4=Scott | first4=D. | last5=Thompson | first5=D.R. | last6=Sagar | first6=P.M. | last7=Moller | first7=H. | last8=Taylor | first8=G.A. | last9=Foley | first9=D.G. | last10=Block | first10=B.A. | last11=Costa | first11=D.P. | date=2006 | title=Migratory shearwaters integrate oceanic resources across the Pacific Ocean in an endless summer | journal=Proceedings of the National Academy of Sciences | volume=103 | issue=34 | pages=12799–12802 | doi=10.1073/pnas.0603715103 | pmid=16908846 | pmc=1568927 | bibcode=2006PNAS..10312799S | doi-access=free }}</ref><ref>{{Cite journal | last1=Carey | first1=M.J. | last2=Phillips | first2=R.A. | last3=Silk | first3=J.R.D. | last4=Shaffer | first4=S.A. | date=2014 | title=Trans-equatorial migration of short-tailed shearwaters revealed by geolocators | journal=Emu | volume=114 | issue=4 | pages=352–359 | doi=10.1071/MU13115| bibcode=2014EmuAO.114..352C | s2cid=84633665 | url=http://nora.nerc.ac.uk/id/eprint/509149/1/MU13115%20Carey%20et%20al%20%20%28REV2%29.doc }}</ref> [[Manx shearwater]]s from the North Atlantic also undertake transequatorial migrations from Western Europe and North America to the waters off Brazil in the South Atlantic.<ref name=j16/> The mechanisms of [[navigation]] are poorly understood, but displacement experiments where individuals were removed from colonies and flown to far-flung release sites have shown that they are able to home in on their colonies with remarkable precision. A Manx shearwater released in [[Boston, Massachusetts|Boston]] returned to its colony in [[Skomer]], [[Wales]] within 13 days, a distance of 5,150 kilometres (3,200 mi).<ref name=j14/> ==Behaviour== ===Food and feeding=== The diet of the procellariids is the most diverse of all the Procellariiformes, as are the methods employed to obtain it. With the exception of the [[giant petrel]]s, all procellariids are exclusively [[ocean|marine]], and the diet of all species is dominated by either fish, [[squid]], [[crustacean]]s and [[carrion]], or some combination thereof.{{sfn|Carboneras|1992a|pp=[https://archive.org/details/handbookofbirdso0001unse/page/223/mode/1up 223–224]}} The majority of species are surface feeders, obtaining food that has been pushed to the surface by other predators or currents, or have floated in death. Among the surface feeders some, principally the gadfly petrels, can obtain food by dipping from flight, while most of the rest feed while sitting on the water. These surface feeders are dependent on their prey being close to the surface, and for this reason procellariids are often found in association with other predators or oceanic convergences. Studies have shown strong associations between many different kinds of [[seabird]]s, including [[wedge-tailed shearwater]]s, and [[dolphin]]s and [[tuna]], which push shoaling fish up towards the surface.<ref name = "AU"/> The gadfly petrels and the Kerguelen petrel mainly feed at night. In so doing they can take advantage of the nocturnal migration of cephalopods and other food species towards the surface.<ref name=Imber1985/><ref name=Harper1987>{{Cite journal | last=Harper | first=Peter C. | date=1987 | title=Feeding behaviour and other notes on 20 species of Procellariiformes at sea | journal=Notornis | volume=34 | issue=3 | pages=169–192 | url=https://notornis.osnz.org.nz/feeding-behaviour-and-other-notes-20-species-procellariiformes-sea | access-date=2022-01-30 | archive-date=2022-01-30 | archive-url=https://web.archive.org/web/20220130121359/https://notornis.osnz.org.nz/feeding-behaviour-and-other-notes-20-species-procellariiformes-sea | url-status=dead }}</ref> The fulmarine petrels are generalists, which for the most part take many species of fish and crustacea. The giant petrels, uniquely for Procellariiformes, will feed on land, eating the carrion of other seabirds and [[pinniped|seals]]. They will also attack the chicks of other seabirds. The diet of the giant petrels varies according to sex, with the females taking more [[krill]] and the males more carrion.<ref name=j20/> All the fulmarine petrels readily feed on fisheries discards at sea, a habit that has been implicated in (but not proved to have caused) the expansion in range of the northern fulmar in the Atlantic.<ref>{{Cite book | last=Thompson | first=P.M. | chapter=Identifying drivers of change: did fisheries play a role in the spread of North Atlantic fulmars? | editor1-last=Boyd | editor1-first=I.L. | editor2-last=Wanless | editor2-first=S. | editor3-last=Camphuysen | editor3-first=C.J. | date=2006 | title=Top Predators in Marine Ecosystems: Their Role in Monitoring and Management | series=Conservation Biology | location=Cambridge, England |publisher=Cambridge University Press | isbn=978-0-521-61256-2 | pages=143–156 | archive-url=https://web.archive.org/web/20060823131229/http://www.abdn.ac.uk/zoology/lighthouse/documents/Thompson_fulmars.pdf | archive-date=2006-08-23 | chapter-url=http://www.abdn.ac.uk/zoology/lighthouse/documents/Thompson_fulmars.pdf}}</ref> [[Image:Broad billed prion.jpg|thumb|The [[broad-billed prion]] (''Pachyptila vittata)'' filters zooplankton from the water with its wide bill. | alt=Photo of a broad-billed prion on land]] The three larger prion species have bills filled with [[Lamellae (zoology)|lamellae]], which act as filters to sift [[zooplankton]] from the water. Water is forced through the lamellae and small prey items are collected. This technique is often used in conjunction with a method known as hydroplaning where the bird dips its bill beneath the surface and propels itself forward with wings and feet as if walking on the water.<ref name=j6/>{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/223/mode/1up 223]}} The diving petrels and many of the shearwaters are proficient divers. While it has long been known that they regularly dive from the surface to pursue prey, using their wings for propulsion,{{sfn|Warham|1990|p=199}} the depth that they are able to dive to was not appreciated (or anticipated) until scientists began to deploy maximum-depth recorders on foraging birds. Studies of both long-distance migrants such as the sooty shearwater and more sedentary species such as the [[black-vented shearwater]] have shown maximum diving depths of {{convert|67|m|ft|abbr=on}} and {{convert|52|m|ft|abbr=on}}.<ref name=j5/><ref>{{Cite journal | last1=Navarro | first1=J. | last2=Votier | first2=S.C. | last3=Phillips | first3=R.A. | date=2014 | title=Diving capabilities of diving petrels | journal=Polar Biology | volume=37 | issue=6 | pages=897–901 | doi=10.1007/s00300-014-1483-0 | doi-access=free | bibcode=2014PoBio..37..897N | hdl=10261/108735 | hdl-access=free }} Supplementary Material.</ref> Tropical shearwaters, such as the wedge-tailed shearwater and the [[Sargasso shearwater]], also dive in order to hunt, making the shearwaters the only tropical seabirds capable of exploiting that ecological niche (all other tropical seabirds feed close to the surface).<ref name=j4/> Many other species of procellariid, from [[white-chinned petrel]]s to [[slender-billed prion]]s, dive to a couple of metres below the surface, though not as proficiently or as frequently as the shearwaters.<ref name=j3/> ===Breeding=== ====Colonies==== [[Image:Kappensturmtaucher.jpg|thumb|The colonies of the [[greater shearwater|great shearwater]] (''Ardenna gravis'') are among the densest of any procellariid, with 1 pair per m<sup>2</sup>. | alt=Photo of a great shearwater in flight ]] The procellariids are colonial, nesting for the most part on islands. These colonies vary in size from over a million birds to just a few pairs, and can be densely concentrated or widely spaced. At one extreme the [[greater shearwater]] nests in concentrations of one pair per square metre in three colonies of more than one million pairs,{{sfn|Brooke|2004|p=28}} whereas the giant petrels nest in clumped but widely spaced territories that barely qualify as colonial. Colonies are usually located near the coast, but some species nest far inland and even at high altitudes. [[Hutton's shearwater]] (''Puffinus huttoni'') breeds in burrows on the sea-facing mountainside of the [[Kaikōura Ranges|Kaikoura Ranges]] on [[South Island]], New Zealand. The colonies are {{cvt|1200|-|1800|m}} above sea level at a distance of {{cvt|12|-|18|km}} from the coast.<ref>{{cite book | year=1990 | chapter=''Puffinus huttoni'' Hutton's Shearwater| editor1-last=Marchant | editor1-first=S. | editor2-last=Higgins | editor2-first=P.G. | title=Handbook of Australian, New Zealand & Antarctic Birds. Volume 1: Ratites to ducks; Part A, Ratites to petrels | location=Melbourne, Victoria | publisher=Oxford University Press | isbn=978-0-19-553068-1 | pages=657–662 | chapter-url=https://nzbirdsonline.org.nz/sites/all/files/111_Hutton's%20Shearwater.pdf }}</ref><ref>{{Cite journal | last1=Sommer | first1=E. | last2=Bell | first2=M. | last3=Bradfield | first3=P. | last4=Dunlop | first4=K. | last5=Gaze | first5=P. | last6=Harrow | first6=G. | last7=McGahan | first7=P. | last8=Morrisey | first8=M. | last9=Walford | first9=D. | last10=Cuthbert | first10=R. | date=2009 | title=Population trends, breeding success and predation rates of Hutton's Shearwater (''Puffinus huttoni''): A 20 year assessment | journal=Notornis | volume=56 | issue=3 | pages=144–153 | url=https://notornis.osnz.org.nz/system/files/Notornis_56_3_144.pdf | access-date=2022-01-22 | archive-date=2022-01-21 | archive-url=https://web.archive.org/web/20220121205539/http://notornis.osnz.org.nz/system/files/Notornis_56_3_144.pdf | url-status=dead }}</ref> Other exceptions are [[Barau's petrel]] (''Pterodroma baraui'') that breeds at {{cvt|2700|m}} on the island of [[Réunion]] in the Indian Ocean,<ref>{{Cite journal | last1=Bretagnolle | first1=Vincent | last2=Attié | first2=Carole | date=1991 | title=Status of Barau's Petrel (''Pterodroma baraui''): colony sites, breeding population and taxonomic affinities | journal=Colonial Waterbirds | volume=14 | issue=1 | pages=25–33 | doi=10.2307/1521275 | jstor=1521275 }}<!-- Table 1 lists high-nesting Procellariiformes--></ref> and the snow petrel (''Pagodroma nivea'') that breeds in Antarctica on mountain ledges up to {{cvt|400|km}} from the open sea.<ref>{{cite book | year=1990 | chapter=''Pagodroma nivea'' Snow Petrel | editor1-last=Marchant | editor1-first=S. | editor2-last=Higgins | editor2-first=P.G. | title=Handbook of Australian, New Zealand & Antarctic Birds. Volume 1: Ratites to ducks; Part A, Ratites to petrels | location=Melbourne, Victoria | publisher=Oxford University Press | isbn=978-0-19-553068-1 | pages=402–410 | chapter-url=https://nzbirdsonline.org.nz/sites/all/files/068_Snow%20Petrel.pdf | access-date=2022-01-22 | archive-date=2022-11-02 | archive-url=https://web.archive.org/web/20221102021333/https://nzbirdsonline.org.nz/sites/all/files/068_Snow%20Petrel.pdf | url-status=dead }}</ref><ref>{{ cite journal | last1=Goldsworthy | first1=P.M. | last2=Thomson | first2=P.G. | date=2000 | title=An extreme inland breeding locality of snow petrels (''Pagodroma nivea'') in the southern Prince Charles Mountains, Antarctica | journal=Polar Biology | volume=23 | issue=10 | pages=717–720 | doi=10.1007/s003000000146 | doi-access= | bibcode=2000PoBio..23..717G | s2cid=21493380 }}</ref> Most seabirds are colonial, and the reasons for colonial behaviour are assumed to be similar, if incompletely understood by scientists. Procellariids for the most part have weak legs and are unable to easily take off, making them highly vulnerable to [[mammal]]ian predators. Most procellariid colonies are located on islands that have historically been free of mammals; for this reason some species cannot help but be colonial as they are limited to a few locations to breed. Even species that breed on continental Antarctica, such as the [[Antarctic petrel]], are forced by habitat preference (snow-free north-facing rock) to breed in just a few locations.{{sfn|Brooke|2004|pp=28–31}} [[Image:Christmas shearwater.jpg|thumb|left|[[Christmas shearwater]]s (''Puffinus nativitatis'') are one of the surface-breeding procellariids. Here a pair engages in some mutual preening. | alt=Photo of a pair of Christmas shearwaters on land under vegetation]] Most procellariids' nests are in burrows or on the surface on open ground, with a smaller number nesting under the cover of vegetation (such as in a forest). All the fulmarine petrels bar the snow petrel nest in the open, the snow petrel instead nesting inside natural crevices. Of the rest of the procellariids the majority nest in burrows or crevices, with a few tropical species nesting in the open. There are several reasons for these differences. The fulmarine petrels are probably precluded from burrowing by their large size (the crevice-nesting snow petrel is the smallest fulmarine petrel) and the high latitudes they breed in, where frozen ground is difficult to burrow into. The smaller size of the other species, and their lack of agility on land, mean that even on islands free from mammal predators they are still vulnerable to [[skua]]s,<ref name=j21/> [[gull]]s and other avian predators, something the aggressive [[stomach oil|oil]]-spitting fulmars are not. The chicks of all species are vulnerable to predation, but the chicks of fulmarine petrels can defend themselves in a similar fashion to their parents. In the higher latitudes there are thermal advantages to burrow nesting, as the temperature is more stable than on the surface, and there is no [[wind-chill]] to contend with. The absence of skuas, gulls and other predatory birds on [[tropical]] islands is why some shearwaters and two species of gadfly petrel ([[Kermadec petrel]] and the [[herald petrel]]) can nest in the open. This has the advantages of reducing competition with burrow nesters from other species and allowing open-ground nesters to nest on [[coral]]line islets without soil for burrowing. Procellariids that burrow in order to avoid predation almost always attend their colonies [[nocturnal animal|nocturnally]] in order to reduce predation as well.<ref name=j22/> Procellariids display high levels of [[philopatry]], exhibiting both natal philopatry and site fidelity. Natal philopatry, the tendency of a bird to breed close to where it hatched, is strong among all the Procellariiformes. The evidence for natal philopatry comes from several sources, not the least of which is the existence of several procellariid species that are endemic to a single island.{{sfn|Brooke|2004|p=111}} The study of [[mitochondrial DNA]] provides evidence of restricted [[gene flow]] between different colonies, and has been used to show philopatry in [[fairy prion]]s.<ref name=j7/> [[Bird ringing]] provides compelling evidence of philopatry; a study of [[Cory's shearwater]]s nesting near [[Corsica]] found that nine out of 61 male chicks that returned to breed at their natal colony actually bred in the burrow they were raised in.<ref name=j8/> This tendency towards philopatry is stronger in some species than others, and several species readily prospect potential new colony sites and colonise them. It is hypothesised that there is a cost to dispersing to a new site, the chance of not finding a mate of the same species, that selects against it for rarer species, whereas there is probably an advantage to dispersal for species that have colony sites that change dramatically during periods of [[Glacial motion|glacial advance or retreat]]. There are differences in the tendency to disperse based on sex, with females being more likely to breed away from the natal site.{{sfn|Brooke|2004|pp=112–113}} ====Mate and site fidelity==== [[Image:Fulmarus glacialis 1 8.jpg|thumb|A [[northern fulmar]] (''Fulmarus glacialis'') pair perform a cackling duet. | alt=Photo of a pair of northern fulmar on a rock]] Procellariids, as well as having strong natal philopatry, exhibit strong site fidelity, returning to the same nesting site, burrow or territory in sequential years. The figure varies for different species but is high for most species, an estimated 91% for [[Bulwer's petrel]]s.<ref name=j9/> The strength of this fidelity can also vary with sex; almost 85% of male [[Cory's shearwater]]s return to the same burrow to breed the year after a successful breeding attempt, while the figure for females is around 76%.<ref name=j10/> This tendency towards using the same site from year to year is matched by strong [[Monogamy in animals|mate fidelity]], with birds breeding with the same partner for many years; it has been suggested that the two are linked, with site fidelity acting as a means in which partnered birds could meet at the beginning of the breeding season.<ref name=j11/> One pair of [[northern fulmar]]s bred as a pair in the same site for 25 years.{{sfn|Carboneras|1992a|pp=[https://archive.org/details/handbookofbirdso0001unse/page/225/mode/1up 225–226]}} Like the [[albatross]]es the procellariids take several years to reach sexual maturity, though due to the greater variety of sizes and lifestyles, the age of first breeding stretches from two or three years in the smaller species to 12 years in the larger ones.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/235/mode/1up 235]}}{{sfn|Carboneras|1992b|p=[https://archive.org/details/handbookofbirdso0001unse/page/275/mode/1up 275]}} The procellariids lack the elaborate breeding dances of the albatrosses, in no small part due to the tendency of most of them to attend colonies at night and breed in burrows, where visual displays are useless. The fulmarine petrels, which nest on the surface and attend their colonies [[Diurnality|diurnally]], do use a repertoire of stereotyped [[ethology|behaviours]] such as cackling, preening, head waving and nibbling, but for most species courtship interactions are limited to some billing (rubbing the two bills together) in the burrow and the vocalisations made by all species. The calls serve a number of functions: they are used territorially to protect burrows or territories and to call for mates. Each call type is unique to a particular species and indeed it is possible for procellariids to identify the sex of the bird calling. It may also be possible to assess the quality of potential mates; a study of [[blue petrel]]s found a link between the [[rhythm]] and duration of calls and the body mass of the bird.<ref name=j23/> The ability of an individual to recognise its mate has been demonstrated in several species.<ref>{{Cite journal | last=Brooke | first=M. de L. | date=1978 | title=Sexual differences in the voice and individual vocal recognition in the Manx shearwater (''Puffinus puffinus'') | journal=Animal Behaviour | volume=26 | pages=622–629 | doi=10.1016/0003-3472(78)90074-X| s2cid=53262540 }}</ref><ref>{{Cite journal | last1=Bretagnolle | first1=Vincent | last2=Lequette | first2=Benoit | date=1990 | title=Structural variation in the call of the Cory's shearwater (''Calonectris diomedea'', Aves, Procellariidae) | journal=Ethology | volume=85 | issue=4 | pages=313–323 | doi=10.1111/j.1439-0310.1990.tb00410.x| bibcode=1990Ethol..85..313B }}</ref> ====Breeding season==== Like most seabirds, the majority of procellariids breed once a year. There are exceptions; many individuals of the larger species, such as the [[white-headed petrel]], will skip a breeding season after successfully [[fledge|fledging]] a chick, and some of the smaller species, such as the [[Christmas shearwater]]s, breed on a nine-month schedule. Among those that breed annually, there is considerable variation as to the timing; some species breed in a fixed season while others breed all year round. [[Climate]] and the availability of food resources are important influences on the timing of procellariid breeding; species that breed at higher [[latitude]]s always breed in the summer as conditions are too harsh in the winter. At lower latitudes many, but not all, species breed continuously. Some species breed seasonally to avoid competition with other species for burrows, to avoid predation or to take advantage of seasonally abundant food. Others, such as the tropical [[wedge-tailed shearwater]], breed seasonally for unknown reasons. Among the species that exhibit seasonal breeding there can be high levels of synchronization, both of time of arrival at the colony and of lay date.{{sfn|Brooke|2004|pp=46–50}} Procellariids begin to attend their nesting colony around one month prior to laying. Males will arrive first and attend the colony more frequently than females, partly in order to protect a site or burrow from potential competitors. Prior to laying there is a period known as the pre-laying exodus in which both the male and female are away from the colony, building up reserves in order to lay and undertake the first incubation stint respectively. This pre-laying exodus can vary in length from 9 days (as in the [[Cape petrel]])<ref name=j24/> to around 50 days in [[Atlantic petrel]]s.<ref name=j25/> All procellariids lay a single white egg per pair per breeding season, in common with the rest of the Procellariiformes. The egg is large compared to that of other birds, weighing 6–24% of the female's weight. Immediately after laying the female goes back to sea to feed while the male takes over incubation. Incubation duties are shared by both sexes in shifts that vary in length between species, individuals and the stage of incubation. The longest recorded shift was 29 days by a [[Murphy's petrel]] from [[Henderson Island (Pitcairn Islands)|Henderson Island]]; the typical length of a [[gadfly petrel]] stint is between 13 and 19 days. Fulmarine petrels, shearwaters and prions tend to have shorter stints, averaging between 3 and 13 days. Incubation takes a long time, from 40 days for the smaller species (such as prions) to around 55 days for the larger species. The incubation period is longer if eggs are abandoned temporarily; procellariid eggs are resistant to chilling and can still hatch after being left unattended for a few days.{{sfn|Carboneras|1992a|p=[https://archive.org/details/handbookofbirdso0001unse/page/227/mode/1up 227]}}{{sfn|Carboneras|1992b|p=[https://archive.org/details/handbookofbirdso0001unse/page/262/mode/1up 262]}} [[Image:Starr 990520-0830 Casuarina equisetifolia.jpg|thumb|The chick of a [[Bonin petrel]] (''Pterodroma hypoleuca'') takes almost three months to fledge. This chick has most of its adult plumage but still retains a considerable amount of down. | alt=Photo of a Bonin petrel chick covered with both feathers and down]] After hatching the chick is brooded by a parent until it is large enough to [[thermoregulation|thermoregulate]] efficiently, and in some cases defend itself from predation. This guard stage lasts a short while for burrow-nesting species (2–3 days) but longer for surface nesting [[fulmar]]s (around 16–20 days) and [[giant petrel]]s (20–30 days). After the guard stage both parents feed the chick. In many species the parent's foraging strategy alternates between short trips lasting 1–3 days and longer trips of 5 days.<ref name=j26/> The shorter trips, which are taken over the continental shelf, benefit the chick with faster growth, but longer trips to more productive [[pelagic zone|pelagic]] feeding grounds are needed for the parents to maintain their own body condition. The meals are composed of both prey items and [[stomach oil]], an [[food energy|energy]]-rich food that is lighter to carry than undigested prey items.<ref name=j27/> This oil is created in a stomach organ known as a [[proventriculus]] from digested prey items, and gives procellariids and other Procellariiformes their distinctive musty smell. Chick development is quite slow for [[bird]]s, with fledging taking place at around two months after hatching for the smaller species and four months for the largest species. The chicks of some species are abandoned by the parents; parents of other species continue to bring food to the nesting site after the chick has left. Chicks put on weight quickly and some can outweigh their parents, although they will slim down before they leave the nest.<ref name="Maynard">{{cite encyclopedia |last=Maynard |first=B.J.|editor1-first=Michael |editor1-last= Hutchins|encyclopedia=Grzimek's Animal Life Encyclopedia |title=Shearwaters, petrels, and fulmars (procellariidae) |edition=2 |year=2003 |publisher=Gale Group|volume=8 Birds I Tinamous and Ratites to Hoatzins |location=Farmington Hills, MI|isbn=978-0-7876-5784-0 |pages=123–127}}</ref> All procellariid chicks fledge by themselves, and there is no further parental care after fledging. Life expectancy of Procellariidae is between 15 and 20 years; the oldest recorded member was a [[northern fulmar]] that was over 50 years.<ref name="Maynard" /> ==Relationship with humans== === Exploitation === [[Image:Puffinus griseus.png|thumb|left|[[Sooty shearwater]]s (''Ardenna grisea'') are still harvested in New Zealand using traditional techniques | alt=Photo of a sooty shearwater in flight ]] Procellariids have been a seasonally abundant source of food for people wherever people have been able to reach their colonies. Early records of human exploitation of shearwaters (along with albatrosses and [[cormorant]]s) come from the remains of [[hunter-gatherer]] [[midden]]s in southern Chile, where sooty shearwaters were taken 5000 years ago.<ref name=j28/> More recently, procellariids have been hunted for food by Europeans, particularly the northern fulmar in Europe, and various species by [[Inuit]],<ref name="Maynard" /> and sailors around the world. The hunting pressure on the [[Bermuda petrel]], or cahow, was so intense that the species nearly became extinct and did go missing for 300 years. The name of one species, the [[providence petrel]], is derived from its (seemingly) miraculous arrival on [[Norfolk Island]], where it provided a windfall for starving European settlers;<ref name = "Anderson"/> within ten years the providence petrel was extinct on Norfolk.<ref name = "Medway"/> Several species of procellariid [[Late Quaternary prehistoric birds#Procellariiformes|have gone extinct]] in the Pacific since the arrival of humans, and their remains have been found in middens dated to that time. More sustainable [[muttonbirding|shearwater harvesting]] industries developed in [[Tasmania]] and New Zealand, where the practice of harvesting what are known as [[shearwater|muttonbird]]s continues today.<ref name="frontiersin.org"/><ref>{{Cite journal | last1=Newman | first1=J. | last2=Scott | first2=D. | last3=Bragg | first3=C. | last4=Mckechnie | first4=S. | last5=Moller | first5=H. | last6=Fletcher | first6=D. | date=2009 | title=Estimating regional population size and annual harvest intensity of the Sooty Shearwater in New Zealand | journal=New Zealand Journal of Zoology | volume=36 | issue=3 | pages=307–323 | doi=10.1080/03014220909510157| s2cid=67827813 | doi-access= }}</ref> ===Threats and conservation=== While some species of procellariid have populations that number in the millions, many species are much less common and several are threatened with [[extinction]]. Human activities have caused dramatic declines in the numbers of some species, particularly species that were originally restricted to one island. According to the [[IUCN]] 43 species are listed as vulnerable or worse, with 12 critically endangered.<ref>{{ cite web | title=Red List: Procellariidae Species | publisher=International Union for Conservation of Nature (IUCN) | url=https://www.iucnredlist.org/search?taxonomies=22672916&searchType=species | access-date=12 February 2022}}</ref> Procellariids are threatened by many threats, but [[introduced species]] on their breeding grounds, [[light pollution]], marine [[fisheries]] particularly bycatch, pollution, exploitation and [[global warming|climate change]] are the main threats measures as the number of species involved.<ref name="frontiersin.org">{{Cite journal | last1=Rodríguez | first1=A. | last2=Arcos | first2=J.M. | last3=Bretagnolle | first3=V. | last4=Dias | first4=M.P. | last5=Holmes | first5=N.D. | last6=Louzao | first6=M. | last7=Provencher | first7=J. | last8=Raine | first8=A.F. | last9=Ramírez | first9=F. | last10=Rodríguez | first10=B. | last11=Ronconi | first11=R.A. | last12=Taylor | first12=R.S. | last13=Bonnaud | first13=E. | last14=Borrelle | first14=S.B. | last15=Cortés | first15=V. | last16=Descamps | first16=S. | last17=Friesen | first17=V.L. | last18=Genovart | first18=M. | last19=Hedd | first19=A. | last20=Hodum | first20=P. | last21=Humphries | first21=G.R.W. | last22=Le Corre | first22=M. | last23=Lebarbenchon | first23=C. | last24=Martin | first24=R. | last25=Melvin | first25=E.F. | last26=Montevecchi | first26=W.A. | last27=Pinet | first27=P. | last28=Pollet | first28=I.L. | last29=Ramos | first29=R. | last30=Russell | first30=J.C. | last31=Ryan | first31=P.G. | last32=Sanz-Aguilar | first32=A. | last33=Spatz | first33=D.R. | last34=Travers | first34=M. | last35=Votier | first35=S.C. | last36=Wanless | first36=R.M. | last37=Woehler | first37=E. | last38=Chiaradia | first38=A. | date=2019 | title=Future directions in conservation research on petrels and shearwaters | journal=Frontiers in Marine Science | volume=6 | pages=94 | doi=10.3389/fmars.2019.00094 | doi-access=free| hdl=2445/139913 | hdl-access=free }}</ref> The most pressing threat for many species, particularly the smaller ones, comes from species introduced to their colonies.<ref name="frontiersin.org"/> Procellariids overwhelmingly breed on islands away from land predators such as mammals, and for the most part have lost the defensive adaptations needed to deal with them (with the exception of the oil-spitting fulmarine petrels). The introduction of mammal predators such as [[feral cat]]s, rats, [[mongoose]]s and mice can have disastrous results for [[island tameness|ecologically naïve]] seabirds.<ref>{{ cite book | last1=Moors | first1=P.J. | last2=Atkinson | first2=I.A.E. | date=1984 | chapter=Alien predation on seabirds | editor1-last=Croxall | editor1-first=J.P. | editor2-last=Evans | editor2-first=P.G.H. | editor3-last=Schreiber | editor3-first=R.W. | title=Status and Conservation of the World's Seabirds | location=Cambridge, UK | publisher=International Council for Bird Preservation | isbn=0-946888-03-5 | pages=667–690 }}</ref> These predators can either directly attack and kill breeding adults, or, more commonly, attack eggs and chicks. Burrowing species that leave their young unattended at a very early stage are particularly vulnerable to attack. Studies on [[grey-faced petrel]]s breeding on New Zealand's [[Whale Island, New Zealand|Whale Island]] (Moutohora) have shown that a population under heavy pressure from [[Norway rat]]s will produce virtually no young during a breeding season, whereas if the rats are controlled (through the use of poison), breeding success is much higher.<ref name ="imber2000"/> That study highlighted the role that non-predatory introduced species can play in harming seabirds; introduced [[rabbit]]s on the island caused little damage to the petrels, other than damaging their burrows, but they acted as a food source for the rats during the non-breeding season, which allowed rat numbers to be higher than they otherwise would be, resulting in more predators for the petrels to contend with. Interactions with introduced species can be quite complex. [[Gould's petrel]]s breed only on two islands, [[Cabbage Tree Island (John Gould Nature Reserve)|Cabbage Tree Island]] and [[Boondelbah Island]] off [[Port Stephens (New South Wales)]]. Introduced rabbits destroyed the forest understory on Cabbage Tree Island; this both increased the vulnerability of the petrels to natural predators and left them vulnerable to the sticky fruits of the birdlime tree (''[[Pisonia umbellifera]]''), 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.<ref name ="car"/> [[Image:Seabirds longlinersm.jpg|thumb|[[Northern fulmars]] (''Fulmarus glacialis'') flocking at a long-lining vessel in the north Pacific]] Larger species of procellariid face similar problems to the albatrosses with [[long-line fishing|long-line fisheries]]. These species readily take offal from fishing boats and will steal bait from the long lines as they are being set, risking becoming snared on the hooks and drowning.<ref name=j29/> In the case of the [[spectacled petrel]] this has led to the species undergoing a large decline and its listing as vulnerable.<ref>{{cite iucn |author=BirdLife International |date=2018 |title=''Procellaria conspicillata'' |volume=2018 |page=e.T22728437A132659002 |doi=10.2305/IUCN.UK.2018-2.RLTS.T22728437A132659002.en |access-date=12 February 2022}}</ref> Diving species, most especially the shearwaters, are also vulnerable to [[gillnet]] fisheries. Studies of gill-net fisheries show that shearwaters (sooty and short-tailed) compose 60% of the seabirds killed by gill-nets in Japanese waters and 40% in Monterey Bay, California in the 1980s,<ref>{{ cite book | last=King | first=W.B. | date=1984 | chapter=Incidental mortality of seabirds in gillnets in the North Pacific | editor1-last=Croxall | editor1-first=J.P. | editor2-last=Evans | editor2-first=P.G.H. | editor3-last=Schreiber | editor3-first=R.W. | title=Status and Conservation of the World's Seabirds | location=Cambridge, UK | publisher=International Council for Bird Preservation | isbn=0-946888-03-5 | pages=709–715 }}</ref> with the total number of shearwaters killed in Japan being between 65,000 and 125,000 per annum over the same study period (1978–1981).<ref>{{ cite book | last=Ogi | first=H. | date=1984 | chapter=Seabird mortality incidental to the Japanese salmon gill-net fishery | editor1-last=Croxall | editor1-first=J.P. | editor2-last=Evans | editor2-first=P.G.H. | editor3-last=Schreiber | editor3-first=R.W. | title=Status and Conservation of the World's Seabirds | location=Cambridge, UK | publisher=International Council for Bird Preservation | isbn=0-946888-03-5 | pages=717–721 }}</ref> Procellariids are vulnerable to other threats as well. Ingestion of plastic [[flotsam]] is a problem for the family as it is for many other seabirds.<ref name=j30/> Once swallowed, this plastic can cause a general decline in the fitness of the bird, or in some cases lodge in the gut and cause a blockage, leading to death by starvation. Procellariids are also vulnerable to general marine pollution, as well as oil spills. Some species, such as the [[Barau's petrel]], the [[Newell's shearwater]] or the [[Cory's shearwater]], which nest high up on large developed islands are victims of light pollution.<ref name=j12/> Chicks that are fledging are attracted to streetlights and are unable to reach the sea. An estimated 20–40% of fledging Barau's petrels are attracted to the streetlights on Réunion.<ref name=j13/> Conservationists are working with governments and fisheries to prevent further declines and increase populations of endangered procellariids. Progress has been made in protecting many colonies where most species are most vulnerable. On 20 June 2001, the [[Agreement on the Conservation of Albatrosses and Petrels]] was signed by seven major fishing nations. The agreement lays out a plan to manage fisheries by-catch, protect breeding sites, promote conservation in the industry, and research threatened species.<ref name="Maynard" /> The developing field of [[island restoration]], where introduced species are removed and native species and habitats restored, has been used in several procellariid recovery programmes.<ref name ="car"/> Invasive species such as rats, feral cats and pigs have been either removed or controlled in many remote islands in the tropical Pacific (such as the [[Northwestern Hawaiian Islands]]), around New Zealand (where island restoration was developed), and in the south Atlantic and [[Indian Ocean]]s. The grey-faced petrels of Whale Island (mentioned above) have been achieving much higher fledging successes after the introduced Norway rats were finally completely removed.<ref name ="imber2000"/> At sea, procellariids threatened by long-line fisheries can be protected using techniques such as setting long-line bait at night, dying the bait blue, setting the bait underwater, increasing the amount of weight on lines and using bird scarers can all reduce the seabird by-catch.<ref>{{ cite report | last1=Brothers | first1=N.P. | last2=Cooper | first2=J. | last3=Løkkeborg | first3=S. | date=1999 | title=The incidental catch of seabirds by longline fisheries: Worldwide review and technical guidelines for mitigation | series= FAO Fisheries Circular No. 937 | location=Rome | publisher=Food and Agriculture Organization of the United Nations | url=https://www.fao.org/3/w9817e/w9817e00.htm }}</ref> The Agreement on the Conservation of Albatrosses and Petrels came into force in 2004 and has been ratified by eight countries, Australia, Ecuador, New Zealand, Spain, South Africa, France, Peru and the United Kingdom. The treaty requires these countries to take specific actions to reduce by-catch and pollution and to remove introduced species from nesting islands.<ref>{{ cite web | title=Agreement on the Conservation of Albatrosses and Petrels (ACAP) | url=https://www.acap.aq/ | publisher=ACAP Secretariat | location=Hobart, Australia |access-date=17 December 2021 }}</ref> ==See also== * [[List of Procellariidae species]] ==References== {{Reflist|refs= <ref name = "Anderson">{{cite journal|author=Anderson, A. |year=1996|title=Origins of Procellariidae Hunting in the Southwest Pacific|journal=International Journal of Osteoarchaeology|volume=6|pages= 403–410|doi=10.1002/(SICI)1099-1212(199609)6:4<403::AID-OA296>3.0.CO;2-0|issue=4}}</ref> <ref name = "AU">{{cite journal|last1=Au |first1=D.W.K. |last2=Pitman |first2=R.L. |year=1986|title= Seabird interactions with dolphins and tuna in the eastern tropical Pacific |journal=Condor|volume=88|pages= 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last=Pennycuick | first=C.J. |year=1982|title=The flight of petrels and albatrosses (Procellariiformes), observed in South Georgia and its vicinity | journal=Philosophical Transactions of the Royal Society of London B | volume=300| pages=75–106 | issue=1098| doi=10.1098/rstb.1982.0158 | jstor=2395926 | bibcode=1982RSPTB.300...75P }}</ref> }} ==Sources== {{refbegin}} *{{ cite book | last=Brooke | first=M. | date=2004 | title=Albatrosses and Petrels Across The World | location=Oxford, UK | publisher=Oxford University Press | isbn=0-19-850125-0 }} *{{ cite book | last=Carboneras | first=C. | year=1992a | chapter=Family Procellariidae (Petrels and Shearwaters) | editor1-last=del Hoyo | editor1-first=J. | editor2-last=Elliott | editor2-first=A. | editor3-last=Sargatal | editor3-first=J. | title=Handbook of the Birds of the World | volume=1: Ostrich to Ducks | location=Barcelona, Spain | publisher=Lynx Edicions | isbn=84-87334-10-5 | pages=216–257 | chapter-url=https://archive.org/details/handbookofbirdso0001unse/page/216/mode/1up | chapter-url-access=registration }} *{{ cite book | last=Carboneras | first=C. | year=1992b | chapter=Family Pelecanoididae (Diving-petrels) | editor1-last=del Hoyo | editor1-first=J. | editor2-last=Elliott | editor2-first=A. | editor3-last=Sargatal | editor3-first=J. | title=Handbook of the Birds of the World | volume=1: Ostrich to Ducks | location=Barcelona, Spain | publisher=Lynx Edicions | isbn=84-87334-10-5 | pages=272–278 | chapter-url=https://archive.org/details/handbookofbirdso0001unse/page/272/mode/1up | chapter-url-access=registration }} *{{ cite book | last=Warham | first=J. | date=1990 | title=The Petrels: Their Ecology and Breeding Systems | location=London | publisher=Academic Press | isbn=0-12-735420-4 }} *{{ cite book | last=Warham | first=J. | date=1996 | title=The Behaviour, Population, Biology and Physiology of the Petrels | location=London | publisher=Academic Press | isbn=0-12-735415-8 }} {{refend}} ==External links== {{Commons category}} {{Wikispecies|Procellariidae}} * [http://www.acap.aq/ The Agreement for the Conservation of Albatrosses and Petrels (ACAP)] Treaty Website * [http://montereybay.com/creagrus/petrels.html Shearwaters and petrels] {{Webarchive|url=https://web.archive.org/web/20081104045230/http://montereybay.com/creagrus/petrels.html |date=2008-11-04 }}: Don Roberson's family page {{Procellariiformes}} {{Birds}} {{Taxonbar|from=Q207767}} {{Authority control}} {{Featured article}} [[Category:Procellariidae| ]] [[Category:Bird families]] [[Category:Seabirds]] [[Category:Extant Rupelian first appearances]] [[Category:Taxa named by William Elford Leach]]
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