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Procellariidae
Cape petrel (Daption capense)
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Procellariiformes
Family: Procellariidae
Leach, 1820
Genera

Macronectes
Fulmarus
Thalassoica
Daption
Pagodroma
Halobaena
Pachyptila
Procellaria
Bulweria
Calonectris
Puffinus
Pelecanoides
Ardenna
Pseudobulweria
Aphrodroma
Pterodroma

Diversity
16 genera, 99 species, of which 3 are extinct

The family Procellariidae is a group of seabirds that comprises the fulmarine petrels, the gadfly petrels, the diving petrels, the prions, and the shearwaters. This family is part of the bird order Procellariiformes (or tubenoses), which also includes the albatrosses and the storm petrels.

The procellariids are the most numerous family of tubenoses, and the most diverse. They range in size from the giant petrels with a wingspan of around 2.0 m (6 ft 7 in), that are almost as large as the albatrosses, to the diving petrels with a wingspan of around 34 cm (13 in) that are similar in size to the little auks 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 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 trans-equatorial migrations. They are colonial breeders, exhibiting long-term mate fidelity and site philopatry. In all species, a single white egg is laid each breeding season. The parents take it in turns to incubate the egg and to forage for food. The feeding area can be at a great distance from the nest site. The incubation times and chick-rearing periods are exceptionally long compared to other birds.

Many procellariids have breeding populations of over several million pairs; others number fewer than 200 birds. Humans have traditionally exploited several species of fulmar 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 fisheries.

Taxonomy and evolution

[edit]

The 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.[1][2] The name is derived from the type genus Procellaria which in turn is derived from the Latin word procella meaning "storm" or "gale".[3] The type genus was named in 1758 by the Swedish naturalist Carl Linnaeus in the tenth edition of his Systema Naturae.[4]

Procellariidae is one of families that make up the order Procellariiformes.[5] Before the introduction of molecular phylogenetics, 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 petrels and Procellariidae containing the petrels, shearwaters and fulmars.[6][7] The family Hydrobatidae was further divided into two subfamilies, the northern storm petrels in Hydrobatinae and the southern or austral storm petrels in Oceanitinae. A 1998 analysis of mitochondrial cytochrome b sequences found there was deep genetic divergence between the two subfamilies.[8] Subsequent large-scale multigene studies found that the two subfamilies were not sister taxa.[9][10][11] The storm petrels were therefore split into two families: Hydrobatidae containing the northern storm petrels and Oceanitidae, containing the southern storm petrels.[5] The multigene genetic studies found that the diving petrels in the family Pelecanoididae were nested within the family Procellariidae.[10][11][12] As a result, the diving petrels was merged into Procellariidae.[5]

The molecular evidence suggests that the albatrosses were the first to diverge from the ancestral stock, and the austral storm petrels next, with the procellariids and northern storm petrels splitting most recently.[10][11][12]

Procellariiformes

Diomedeidae – albatrosses (21 species)

Oceanitidae – austral storm petrels (9 species)

Hydrobatidae – northern storm petrels (18 species)

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.[13] The genus was therefore split and a group of species moved into the resurrected genus Ardenna.[5][14] The other genera within the family were found to be monotypic but the relationships between the genera remained unclear.[13] This changed when a multigene genetic study published in 2021 provided a genus-level phylogeny of the family.[15]

Procellariidae

Pagodroma – snow petrel

Thalassoica – Antarctic petrel

Daption – Cape petrel

Macronectes – giant petrels (2 species)

Fulmarus – fulmars (2 species)

Pelecanoides – diving petrels (4 species)

Halobaena – blue petrel

Pachyptila – prions (7 species)

Aphrodroma – Kerguelen petrel

Pterodroma – gadfly petrels (35 species)

Bulweria – petrels (2 extant species)

Pseudobulweria – petrels (4 species)

Procellaria – petrels (5 species)

Puffinus – shearwaters (21 species)

Calonectris – shearwaters (4 species)

Ardenna – shearwaters (7 species)

There are 99 species of procellariid in 16 genera.[5] The family has usually been broken up into four fairly distinct groups; the fulmarine petrels, the gadfly petrels, the prions, and the shearwaters. With the inclusion of the diving petrels there are now five main groups.[16][17]

  • The fulmarine petrels include the largest procellariids, the giant petrels, 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 morphologically by their skull features, particularly the long prominent nasal tubes.[18]
  • The four diving petrels are the smallest procellariids with lengths of around 20 cm (7.9 in) and wingspans of 33 cm (13 in). 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.[19]
  • 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, 26–46 cm (10–18 in) in length, and are long winged with short hooked bills.[20] They are most closely related to Kerguelen petrel which is placed in its own genus Aphrodroma.[15]
  • 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 that they use to filter plankton somewhat as baleen whales 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, 25–30 cm (9.8–11.8 in) in length, with a prominent dark M-shaped mark across the upperwing of their grey plumage. All are restricted to the southern hemisphere.[21]
  • The shearwaters are adapted for diving after prey instead of foraging on the ocean's surface; several species have been recorded diving deeper than 30 m (98 ft).[22] They are known for the long trans-equatorial migrations undertaken by many species.[23] 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.[5]

Morphology and flight

[edit]
Photo of a giant petrel in flight
The flight of giant petrels is aided by a shoulder-lock that holds their wing out without effort.
Dark brown bird with outstretched wings prepares to take off from sandy beach
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 185 to 205 cm (73 to 81 in), is almost as large as albatrosses; the smallest, the diving petrels have a wingspan of 30 to 38 cm (12 to 15 in) and are similar in size to little auks or dovekies in the family Alcidae.[24][25][26] There are no obvious differences between the sexes, although females tend to be slighter.[27][28] Like all Procellariiformes, the procellariids have a characteristic tubular nasal passage used for olfaction.[29] This ability to smell helps to locate patchily distributed prey at sea and may help locate nesting colonies. The plumage of the procellariids is usually dull, with greys, bluish greys, blacks and browns being the usual colours,[18] although some species have striking patterns such as the Cape petrel.[30]

The technique of 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.[31] 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.[32] Because of the high speeds required for flight, procellariids need to either run or face into a strong wind in order to take off.[33]

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.[31] Gadfly petrels often feed on the wing, snapping prey without landing on the water. The flight of the smaller prions is similar to that of the storm petrels, being highly erratic and involving weaving and even looping the loop. The wings of all species are long and stiff. In some species of shearwater the wings are used to power the birds underwater while diving for prey. Their heavier wing loadings, in comparison with surface-feeding procellariids, allow these shearwaters to achieve considerable depths (below 70 m (230 ft) in the case of the short-tailed shearwater).[34]

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.[35] The exceptions to this are the two species of giant petrel, which have strong legs used when they feed on land.[18]

Distribution and migration

[edit]
Photo of a flock of shearwaters in flight
Million-strong flocks of shearwaters migrate from New Zealand to Alaska every year.

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.[36][37] Among the groups, the fulmarine petrels have a mostly polar distribution, with most species living around Antarctica and one, the northern fulmar ranging in the Northern Atlantic and Pacific Oceans.[23] 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.[38] 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.[23]

Many procellariids undertake long annual 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.[39][40] Manx shearwaters from the North Atlantic also undertake transequatorial migrations from Western Europe and North America to the waters off Brazil in the South Atlantic.[41] 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 returned to its colony in Skomer, Wales within 13 days, a distance of 5,150 kilometres (3,200 mi).[42]

Behaviour

[edit]

Food and feeding

[edit]

The diet of the procellariids is the most diverse of all the Procellariiformes, as are the methods employed to obtain it. With the exception of the giant petrels, all procellariids are exclusively marine, and the diet of all species is dominated by either fish, squid, crustaceans and carrion, or some combination thereof.[43]

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 seabirds, including wedge-tailed shearwaters, and dolphins and tuna, which push shoaling fish up towards the surface.[44] 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.[20][45]

The fulmarine petrels are generalists, which for the most part take many species of fish and crustacea. The giant petrels, uniquely for Procellariiformes, will feed on land, eating the carrion of other seabirds and seals. They will also attack the chicks of other seabirds. The diet of the giant petrels varies according to sex, with the females taking more krill and the males more carrion.[46] 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.[47]

Photo of a broad-billed prion on land
The broad-billed prion (Pachyptila vittata) filters zooplankton from the water with its wide bill.

The three larger prion species have bills filled with 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.[48][49]

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,[50] 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 67 m (220 ft) and 52 m (171 ft).[51][52] 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).[53] Many other species of procellariid, from white-chinned petrels to slender-billed prions, dive to a couple of metres below the surface, though not as proficiently or as frequently as the shearwaters.[54]

Breeding

[edit]

Colonies

[edit]
Photo of a great shearwater in flight
The colonies of the great shearwater (Ardenna gravis) are among the densest of any procellariid, with 1 pair per m2.

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,[55] whereas the giant petrels nest in clumped but widely spaced territories that barely qualify as colonial. Colonies are usually located near the coast, but some species nest far inland and even at high altitudes. Hutton's shearwater (Puffinus huttoni) breeds in burrows on the sea-facing mountainside of the Kaikoura Ranges on South Island, New Zealand. The colonies are 1,200–1,800 m (3,900–5,900 ft) above sea level at a distance of 12–18 km (7.5–11.2 mi) from the coast.[56][57] Other exceptions are Barau's petrel (Pterodroma baraui) that breeds at 2,700 m (8,900 ft) on the island of Réunion in the Indian Ocean,[58] and the snow petrel (Pagodroma nivea) that breeds in Antarctica on mountain ledges up to 400 km (250 mi) from the open sea.[59][60]

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 mammalian 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.[61]

Photo of a pair of Christmas shearwaters on land under vegetation
Christmas shearwaters (Puffinus nativitatis) are one of the surface-breeding procellariids. Here a pair engages in some mutual preening.

Most procellariids' nests are in burrows or on the surface on open ground, with a smaller number nesting under the cover of vegetation (such as in a forest). All the fulmarine petrels bar the snow petrel nest in the open, the snow petrel instead nesting inside natural crevices. Of the rest of the procellariids the majority nest in burrows or crevices, with a few tropical species nesting in the open. There are several reasons for these differences. The fulmarine petrels are probably precluded from burrowing by their large size (the crevice-nesting snow petrel is the smallest fulmarine petrel) and the high latitudes they breed in, where frozen ground is difficult to burrow into. The smaller size of the other species, and their lack of agility on land, mean that even on islands free from mammal predators they are still vulnerable to skuas,[62] gulls and other avian predators, something the aggressive oil-spitting fulmars are not. The chicks of all species are vulnerable to predation, but the chicks of fulmarine petrels can defend themselves in a similar fashion to their parents. In the higher latitudes there are thermal advantages to burrow nesting, as the temperature is more stable than on the surface, and there is no wind-chill to contend with. The absence of skuas, gulls and other predatory birds on tropical 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 coralline islets without soil for burrowing. Procellariids that burrow in order to avoid predation almost always attend their colonies nocturnally in order to reduce predation as well.[63]

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.[64] The study of mitochondrial DNA provides evidence of restricted gene flow between different colonies, and has been used to show philopatry in fairy prions.[65] Bird ringing provides compelling evidence of philopatry; a study of Cory's shearwaters 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.[66] This tendency towards philopatry is stronger in some species than others, and several species readily prospect potential new colony sites and colonise them. It is hypothesised that there is a cost to dispersing to a new site, the chance of not finding a mate of the same species, that selects against it for rarer species, whereas there is probably an advantage to dispersal for species that have colony sites that change dramatically during periods of glacial advance or retreat. There are differences in the tendency to disperse based on sex, with females being more likely to breed away from the natal site.[67]

Mate and site fidelity

[edit]
Photo of a pair of northern fulmar on a rock
A northern fulmar (Fulmarus glacialis) pair perform a cackling duet.

Procellariids, as well as having strong natal philopatry, exhibit strong site fidelity, returning to the same nesting site, burrow or territory in sequential years. The figure varies for different species but is high for most species, an estimated 91% for Bulwer's petrels.[68] The strength of this fidelity can also vary with sex; almost 85% of male Cory's shearwaters return to the same burrow to breed the year after a successful breeding attempt, while the figure for females is around 76%.[69] This tendency towards using the same site from year to year is matched by strong mate fidelity, with birds breeding with the same partner for many years; it has been suggested that the two are linked, with site fidelity acting as a means in which partnered birds could meet at the beginning of the breeding season.[70] One pair of northern fulmars bred as a pair in the same site for 25 years.[71] Like the albatrosses 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.[24][72]

The procellariids lack the elaborate breeding dances of the albatrosses, in no small part due to the tendency of most of them to attend colonies at night and breed in burrows, where visual displays are useless. The fulmarine petrels, which nest on the surface and attend their colonies diurnally, do use a repertoire of stereotyped behaviours such as cackling, preening, head waving and nibbling, but for most species courtship interactions are limited to some billing (rubbing the two bills together) in the burrow and the vocalisations made by all species. The calls serve a number of functions: they are used territorially to protect burrows or territories and to call for mates. Each call type is unique to a particular species and indeed it is possible for procellariids to identify the sex of the bird calling. It may also be possible to assess the quality of potential mates; a study of blue petrels found a link between the rhythm and duration of calls and the body mass of the bird.[73] The ability of an individual to recognise its mate has been demonstrated in several species.[74][75]

Breeding season

[edit]

Like most seabirds, the majority of procellariids breed once a year. There are exceptions; many individuals of the larger species, such as the white-headed petrel, will skip a breeding season after successfully fledging a chick, and some of the smaller species, such as the Christmas shearwaters, breed on a nine-month schedule. Among those that breed annually, there is considerable variation as to the timing; some species breed in a fixed season while others breed all year round. Climate and the availability of food resources are important influences on the timing of procellariid breeding; species that breed at higher latitudes 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.[76]

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)[77] to around 50 days in Atlantic petrels.[78] All procellariids lay a single white egg per pair per breeding season, in common with the rest of the Procellariiformes. The egg is large compared to that of other birds, weighing 6–24% of the female's weight. Immediately after laying the female goes back to sea to feed while the male takes over incubation. Incubation duties are shared by both sexes in shifts that vary in length between species, individuals and the stage of incubation. The longest recorded shift was 29 days by a Murphy's petrel from Henderson Island; the typical length of a gadfly petrel 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.[79][80]

Photo of a Bonin petrel chick covered with both feathers and down
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.

After hatching the chick is brooded by a parent until it is large enough to thermoregulate efficiently, and in some cases defend itself from predation. This guard stage lasts a short while for burrow-nesting species (2–3 days) but longer for surface nesting fulmars (around 16–20 days) and giant petrels (20–30 days). After the guard stage both parents feed the chick. In many species the parent's foraging strategy alternates between short trips lasting 1–3 days and longer trips of 5 days.[81] The shorter trips, which are taken over the continental shelf, benefit the chick with faster growth, but longer trips to more productive 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 energy-rich food that is lighter to carry than undigested prey items.[82] 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 birds, 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.[83] 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.[83]

Relationship with humans

[edit]

Exploitation

[edit]
Photo of a sooty shearwater in flight
Sooty shearwaters (Ardenna grisea) are still harvested in New Zealand using traditional techniques

Procellariids have been a seasonally abundant source of food for people wherever people have been able to reach their colonies. Early records of human exploitation of shearwaters (along with albatrosses and cormorants) come from the remains of hunter-gatherer middens in southern Chile, where sooty shearwaters were taken 5000 years ago.[84] More recently, procellariids have been hunted for food by Europeans, particularly the northern fulmar in Europe, and various species by Inuit,[83] 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;[85] within ten years the providence petrel was extinct on Norfolk.[86] Several species of procellariid have gone extinct in the Pacific since the arrival of humans, and their remains have been found in middens dated to that time. More sustainable shearwater harvesting industries developed in Tasmania and New Zealand, where the practice of harvesting what are known as muttonbirds continues today.[87][88]

Threats and conservation

[edit]

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.[89] Procellariids are threatened by many threats, but introduced species on their breeding grounds, light pollution, marine fisheries particularly bycatch, pollution, exploitation and climate change are the main threats measures as the number of species involved.[87]

The most pressing threat for many species, particularly the smaller ones, comes from species introduced to their colonies.[87] 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 cats, rats, mongooses and mice can have disastrous results for ecologically naïve seabirds.[90] 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 petrels breeding on New Zealand's Whale Island (Moutohora) have shown that a population under heavy pressure from Norway rats will produce virtually no young during a breeding season, whereas if the rats are controlled (through the use of poison), breeding success is much higher.[91] That study highlighted the role that non-predatory introduced species can play in harming seabirds; introduced rabbits on the island caused little damage to the petrels, other than damaging their burrows, but they acted as a food source for the rats during the non-breeding season, which allowed rat numbers to be higher than they otherwise would be, resulting in more predators for the petrels to contend with. Interactions with introduced species can be quite complex. Gould's petrels breed only on two islands, Cabbage Tree Island and Boondelbah Island off Port Stephens (New South Wales). Introduced rabbits destroyed the forest understory on Cabbage Tree Island; this both increased the vulnerability of the petrels to natural predators and left them vulnerable to the sticky fruits of the birdlime tree (Pisonia umbellifera), 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.[92]

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 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.[93] In the case of the spectacled petrel this has led to the species undergoing a large decline and its listing as vulnerable.[94] 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,[95] 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).[96]

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.[97] 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.[98] 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.[99]

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.[83] 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.[92] 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 Oceans. 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.[91] 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.[100] 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.[101]

See also

[edit]

References

[edit]

Sources

[edit]
  • Brooke, M. (2004). Albatrosses and Petrels Across The World. Oxford, UK: Oxford University Press. ISBN 0-19-850125-0.
  • Carboneras, C. (1992a). "Family Procellariidae (Petrels and Shearwaters)". In del Hoyo, J.; Elliott, A.; Sargatal, J. (eds.). Handbook of the Birds of the World. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. pp. 216–257. ISBN 84-87334-10-5.
  • Carboneras, C. (1992b). "Family Pelecanoididae (Diving-petrels)". In del Hoyo, J.; Elliott, A.; Sargatal, J. (eds.). Handbook of the Birds of the World. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. pp. 272–278. ISBN 84-87334-10-5.
  • Warham, J. (1990). The Petrels: Their Ecology and Breeding Systems. London: Academic Press. ISBN 0-12-735420-4.
  • Warham, J. (1996). The Behaviour, Population, Biology and Physiology of the Petrels. London: Academic Press. ISBN 0-12-735415-8.
[edit]
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from Grokipedia
Procellariidae is a family of tube-nosed seabirds belonging to the order Procellariiformes, encompassing fulmars, gadfly petrels, prions, and shearwaters as its primary subgroups.[1][2] This diverse assemblage represents the largest family within Procellariiformes, with over 80 species distributed across 12 to 16 genera, inhabiting oceans worldwide but concentrated in temperate and subantarctic waters.[3][4] These birds exhibit specialized adaptations for a pelagic existence, including elongated wings suited for dynamic soaring over wave fronts and tubular nostrils that facilitate salt excretion via supraorbital glands and enhance olfactory detection of prey such as cephalopods, fish, and crustaceans from considerable distances.[5][6] Foraging occurs primarily at the sea surface, with some species employing pursuit plunging or scavenging behaviors, while breeding is confined to remote colonies on islands or coastal cliffs, often involving burrow or crevice nests and protracted incubation periods exceeding 50 days for a single egg.[2][5] Procellariids undertake extensive migrations, with certain shearwaters traversing hemispheres between breeding sites and wintering grounds, underscoring their reliance on ocean currents and wind patterns.[3] Defining characteristics include sexual size dimorphism in some taxa and vocal or aerial displays during courtship, yet the family faces significant anthropogenic pressures, notably bycatch in longline fisheries and habitat degradation from invasive predators at nesting locales, contributing to population declines in multiple species.[7][6]

Taxonomy and Systematics

Phylogenetic Relationships and Classification

The family Procellariidae, encompassing petrels, shearwaters, and fulmars, forms one of four families within the monophyletic order Procellariiformes, alongside Diomedeidae (albatrosses) and the two storm-petrel families Oceanitidae and Hydrobatidae.[8] It includes approximately 99 extant species distributed across 16 genera, such as Ardenna (larger shearwaters), Calonectris (gansu shearwaters), Macronectes (giant petrels), Puffinus (smaller shearwaters), Pterodroma (gadfly petrels), and Fulmarus (fulmars).[9] [10] Molecular phylogenetic studies, employing mitochondrial markers like cytochrome b and nuclear genes, have robustly confirmed the monophyly of Procellariidae within Procellariiformes, with internal divergences reflecting biogeographic patterns and ecological specializations.[11] For instance, analyses of shearwater genera reveal distinct clades corresponding to body size variations and oceanic distributions, such as the separation of larger, long-distance migrant forms in Ardenna from smaller, tropical Puffinus species, driven by adaptive radiations in different hemispheres.[12] Genome-wide approaches further demonstrate that substitution rate heterogeneity across lineages does not undermine these topologies, supporting relaxed clock models for estimating divergence times among procellariid subgroups.[13] Taxonomic classification within Procellariidae remains dynamic, with debates centering on species boundaries informed by integrative evidence versus traditional morphology. Proposals for splitting complexes, such as in Pterodroma petrels and Puffinus shearwaters, rely on genetic divergence, vocalization differences, and subtle plumage variations, as seen in recent elevations of subspecies to full species status based on mitochondrial and nuclear data; these contrast with lumping advocated by morphologists emphasizing intergradation in breeding grounds.[14] Recent fossil validations, including the Pliocene Ardenna buchananbrowni—the earliest known diving member of its genus—bolster the delineation of modern Ardenna taxa by confirming ancestral specializations for underwater pursuit foraging, aligning genetic clades with osteological traits.[15] Such integrations highlight how molecular and paleontological data refine classifications, though consensus lags for vocally distinct island endemics pending comprehensive sampling.

Fossil Record and Evolutionary History

The fossil record of Procellariidae documents the family's origins in the Paleogene, with the earliest confirmed remains dating to the Early Oligocene.[16] Subsequent diversification is evidenced by Miocene fossils, including a new petrel species from the early Miocene Gaiman Formation in Patagonia, Argentina, represented by a carpometacarpus indicating early adaptation to marine environments.[17] Mid-Miocene records from California further attest to the presence of fulmarine petrels, the sister group to modern Macronectes, suggesting initial radiation into scavenging niches amid expanding oceanic habitats.[18] Pliocene fossils reveal increased morphological specialization, such as diving adaptations in shearwaters. A recently described species, Ardenna buchananbrowni, from Pliocene deposits in Taranaki, New Zealand (approximately 3.5–5 million years ago), represents the smallest and oldest known diving-specialized member of the genus Ardenna, with skeletal features like a robust humerus enabling pursuit diving for prey; this find predates molecular estimates of the Puffinus-Ardenna divergence by over 10 million years, highlighting underrepresentation in prior fossil data.[15] Similarly, Macronectes tinae, the first fossil giant petrel from the Pliocene Tangahoe Formation in New Zealand (circa 3 million years ago), exhibits a smaller body size than extant congeners but a massive, hooked bill suited for carcass dismemberment, indicating early evolution of aggressive scavenging behaviors tied to marine productivity.[18] Evolutionary trends in Procellariidae reflect adaptations to pelagic niches, including enhanced olfaction via tubular nostrils and efficient gliding flight, as inferred from consistent skeletal proportions across Miocene-to-Pleistocene fossils.[19] Shearwater body sizes varied with prey availability and island isolation, while the family's persistence through Pleistocene volcanism on breeding islands demonstrates resilience to episodic natural disturbances, unlike the selective pressures from modern human activities.[20] Overall, fossil evidence links diversification to Neogene oceanic cooling and upwelling intensification, which boosted krill and squid populations central to procellariid diets, though direct causal chains remain constrained by sparse pre-Miocene records.[21]

Morphology and Physiology

Physical Characteristics

Members of the Procellariidae exhibit a characteristic bill structure consisting of multiple horny plates, a straight and deeply grooved culmen, a hooked tip, and paired tubular nostrils extending along the dorsal midline.[22] These nostrils are enclosed in keratinized tubes, a diagnostic feature of the family derived from dissections and morphological examinations.[22] The bills vary in robustness across species, with larger forms like giant petrels (Macronectes spp.) possessing heavier, more massive structures adapted to their predatory habits, as observed in specimen measurements.[14] Body size shows considerable variation, ranging from smaller species such as prions (Pachyptila spp.) with lengths of approximately 23–28 cm and body masses around 120–170 g to the largest, southern giant petrels (Macronectes giganteus), where males reach lengths up to 81–99 cm, wingspans of 150–210 cm, and masses up to 5 kg.[23] Wingspans in the family generally span 40–210 cm, with longer wings in shearwaters and petrels facilitating extended pelagic flight, based on field measurements and museum specimens.[23] Sexual dimorphism in size is minimal in most species, with males and females exhibiting similar plumage and dimensions, though pronounced in Macronectes where males exceed females by up to 40% in mass.[24] [25] Plumage is typically dense and soft, featuring subdued tones of black, white, gray, and brown for visual distinction among species, as documented in taxonomic descriptions.[26] Patterns vary, including underwing markings and mottled upperparts, but lack sexual or seasonal differences in most cases.[26] Physiologically, procellariids possess well-developed supraorbital salt glands above the eyes, enabling excretion of concentrated saline solutions, confirmed through anatomical dissections and osmoregulatory studies in related procellariiforms.[27] These glands, larger in marine-adapted species, maintain ionic balance via active transport mechanisms observed in experimental assays.[28]

Adaptations for Flight, Olfaction, and Marine Life

Procellariids are adapted for sustained pelagic flight through dynamic soaring, a biomechanical strategy that leverages vertical wind gradients over the ocean to minimize powered flapping. This involves repeated cycles of ascent in headwinds and descent in tailwinds, converting potential energy from wind shear into forward momentum. Empirical data from biologging devices reveal ground speeds up to 28 m/s (100 km/h) in species such as shearwaters and petrels, with energy expenditure approaching zero during optimal conditions due to the passive harnessing of atmospheric shear forces.[29][30] Their olfactory system represents a key sensory adaptation for open-ocean navigation and prey location, surpassing visual reliance in low-light or foggy conditions common at sea. The tubular nostrils characteristic of Procellariiformes funnel air into enlarged olfactory bulbs, enabling detection of dilute plankton-derived odors like dimethyl sulfide from kilometers away. Field experiments demonstrate species-specific attraction to krill-associated volatiles, such as 3-methylpyrazine, prompting foraging approaches; for example, Antarctic petrels and prions home in on these scents, confirming olfaction's causal role in locating ephemeral food patches.[31][32][33] Physiological traits support marine existence via modulated buoyancy and pursuit diving, though capabilities vary phylogenetically within the family. Shearwaters achieve depths exceeding 90 m using wing-powered propulsion, facilitated by robust pectoral musculature rich in slow-twitch fibers for aerobic endurance and flattened humeri reducing drag. In contrast, prions rely on shallower foot-propelled dives, with higher overall buoyancy from pneumatic skeletons aiding surface recovery. These differences correlate with bone wall thickness and muscle myoglobin levels, enhancing oxygen delivery and streamlining for brief submersion while preserving flight efficiency upon resurfacing.[34][35][15]

Distribution and Habitat

Global Range and Habitat Preferences

Procellariidae species inhabit all major oceans, spanning tropical to polar latitudes, with breeding concentrated on oceanic islands rather than continental landmasses. At sea, they occupy pelagic environments, often associating with productive marine areas such as upwelling zones that enhance prey availability through nutrient upwelling.[22][36] Empirical at-sea surveys document their widespread non-breeding distributions, revealing concentrations in temperate and subpolar waters where oceanographic productivity supports foraging.[37] Breeding occurs predominantly in the Southern Hemisphere, with highest species richness on sub-Antarctic islands in the southern Indian and Atlantic Oceans, including sites like South Georgia and the Antarctic Peninsula.[36][14] Certain genera exhibit Northern Hemisphere breeding, such as Calonectris shearwaters on North Atlantic islands like the Azores and in the Mediterranean.[38] Core breeding ranges remain stable on predator-free remote islands, where banding and GPS data indicate limited shifts despite tracking individual movements.[39] Habitat preferences favor isolated islands with suitable burrowing substrates, vegetation cover for shade, and inaccessibility to terrestrial predators, leading to nest sites in soil cavities or under rocks.[8] These selections minimize mammalian predation risks, as evidenced by higher burrow densities on islands lacking introduced predators.[40] Oceanic dependencies are underscored by their reliance on surrounding marine productivity, with distributions correlating to areas of elevated chlorophyll concentrations from satellite and survey data.[36]

Migration Patterns and Dispersal

Species within Procellariidae exhibit diverse migration strategies, ranging from extensive trans-equatorial journeys to localized post-breeding dispersal, primarily driven by wind regimes, ocean currents, and prey distributions revealed through geolocation and satellite tracking. Many shearwaters and petrels undertake long-distance migrations that exploit productive marine fronts, with routes often following prevailing winds to minimize energetic costs.[41][42] Sooty shearwaters (Ardenna grisea) exemplify extreme migratory behavior, completing annual roundtrips of approximately 64,000 km in a figure-eight trajectory across the Pacific Ocean, integrating resources from temperate and subarctic waters while avoiding oligotrophic zones.[43] Tracking data indicate fidelity to dynamic oceanographic features, such as upwelling systems, with individuals covering over 1,000 km per day under favorable conditions.[44] Similar trans-equatorial patterns occur in other procellariids, including Cory's shearwaters (Calonectris diomedea), which migrate southward to the Benguela Current off southern Africa, leveraging nutrient-rich upwellings for foraging.[45] Juvenile dispersal often contrasts with adult patterns, featuring wider ranging to prospect for future breeding sites or resources, influenced by wind patterns and prey patches, whereas adults display higher site fidelity during non-breeding periods.[46] In giant petrels (Macronectes spp.), adults exhibit resident-like behavior near colonies post-breeding, concentrating foraging in shelf-break and middle-shelf waters, while juveniles explore broader areas, with sex-specific differences showing females dispersing farther.[47][48] Genetic analyses underscore strong natal philopatry across Procellariidae, where individuals return to birth colonies for breeding, fostering fine-scale population differentiation despite migratory phases that could facilitate gene flow; this is evidenced by low inter-colony dispersal rates and isolation by distance in species like shearwaters and prions.[49][50] Such philopatry, combined with environmental cues like consistent current systems, maintains route fidelity across generations.[51]

Behavior and Ecology

Foraging Strategies and Diet

Procellariids primarily employ surface-seizing and shallow-plunging techniques to capture prey while in flight or on the water surface, targeting epipelagic and mesopelagic organisms such as cephalopods, myctophid fish, and crustaceans including krill and amphipods.[52][53] Regurgitate analyses and DNA metabarcoding of fecal samples consistently reveal these marine invertebrates and fish as dominant components, with cephalopods comprising 25-57% by mass in species like the white-chinned petrel (Procellaria aequinoctialis), alongside comparable proportions of fish and crustaceans.[54][55] Stable isotope ratios (δ¹³C and δ¹⁵N) from tissues such as feathers and eggs corroborate this, indicating trophic levels around 3-4, with δ¹⁵N values reflecting krill-based diets shifting to higher piscivory during chick-rearing in species like Wilson's storm-petrel (Oceanites oceanicus).[56] Olfaction plays a central role in locating prey patches across vast foraging ranges extending thousands of kilometers, enabling detection of volatile compounds like dimethyl sulfide (DMS) produced by phytoplankton and grazing zooplankton at concentrations as low as 10⁻¹² mol l⁻¹.[31] Procellariids such as prions, storm-petrels, and gadfly petrels use upwind zigzag flight patterns to map an olfactory landscape tied to oceanographic features like fronts and upwellings, transitioning to area-restricted searches upon encountering scent plumes.[31] This sensory reliance surpasses visual cues for initial prey detection, particularly at night when many target bioluminescent squid, though visual confirmation aids final capture.[31] Diel and seasonal variations influence strategies, with empirical tracking data showing nocturnal dives for vertically migrating prey and diurnal surface feeding on krill aggregations during productive seasons.[52] Larger taxa like giant petrels (Macronectes spp.) diverge by incorporating scavenging of marine mammal and penguin carcasses—up to dominant portions in Antarctic colonies—and kleptoparasitism on other seabirds, as observed in interactions with albatrosses.[57][58] Stable isotope signatures in these species reflect opportunistic shifts to higher trophic carrion, contrasting the consistent mid-trophic marine diets of smaller procellariids.[56]

Social Structure and Communication

Members of the Procellariidae family breed in colonies, often comprising thousands to millions of pairs, but with burrows or nests spaced several meters apart to minimize physical interactions and aggression between neighbors.[59] This spacing results in relatively loose colonial structures compared to more densely packed seabird groups, with aggression primarily limited to occasional disputes over shared burrow sites or resources.[60] Long-term monogamous pair bonds, typically lasting multiple seasons, are maintained through mutual vocal duets and calls that encode individual signatures, enabling mate recognition and coordination during breeding activities such as incubation shifts.[61] For instance, in Manx shearwaters (Puffinus puffinus), playback experiments demonstrate that these vocalizations facilitate pair re-establishment upon return to colonies, with calls interspersed between aerial and burrow contexts to reinforce bonds.[62][63] Olfactory cues complement acoustic signals in social recognition, particularly for locating partners and burrows in dark, burrow-nesting environments. Antarctic prions (Pachyptila desolata) discriminate mates via chemical signals, reducing reliance on potentially predator-attracting vocalizations.[64] At sea, procellariids are predominantly solitary foragers, but some species form loose aggregations during prey encounters, leveraging social cues and olfaction to detect food patches such as those emitting dimethyl sulfide from plankton.[31] In scavenging guilds, like those involving giant petrels (Macronectes spp.), temporary dominance hierarchies emerge at carcasses, where larger males assert priority access over females and subordinates through displays and physical contests.[65] Vocal repertoires vary by genus but generally include guttural croaks, drones, and trills for territory defense and inter-pair signaling, with empirical recordings showing species-specific patterns that minimize overlap in noisy colonies.[14] These communication strategies adapt to the family's pelagic lifestyle, balancing individual foraging efficiency with breeding-season social needs.[66]

Reproduction and Life Cycle

Members of the Procellariidae family typically breed annually, though larger species may breed biennially, laying a single egg per clutch in burrows or on open ground.[9] Both parents share incubation duties, with periods ranging from 50 to 60 days on average, varying by species size and environmental conditions.[67] Incubation shifts last several days to weeks, allowing parents to forage at sea for stomach oil-rich prey to sustain the breeding effort.[9] Upon hatching, chicks are brooded initially by one parent while the other forages, transitioning to longer absences as the chick develops thermoregulation and waterproofing.[8] Chick-rearing durations span 2 to 6 months, with growth rates empirically linked to parental provisioning success and oceanographic food availability; larger species exhibit prolonged nestling phases up to 7 months in some cases.[8] Fledglings depart independently to sea, often without parental guidance, marking the end of the breeding cycle.[68] Sexual maturity is deferred, with first breeding occurring between 4 and 10 years of age, reflecting slow life histories optimized for high adult survival over rapid reproduction.[69] Banding studies reveal maximum lifespans exceeding 50 years in many species, contrasting with low annual fecundity of approximately 0.5 to 1 fledgling per pair, embodying a classic trade-off where extended longevity compensates for minimal reproductive output.[70] This K-selected strategy prioritizes survival and mate fidelity, with empirical demographic models showing that adult mortality rates below 10% annually underpin population persistence despite sporadic breeding failures.[71]

Population Dynamics and Threats

Monitored seabird populations worldwide, including those of Procellariiformes, declined by 69.7% from 1950 to 2010, based on a database of over 5,000 records from long-term colony monitoring.[72] Procellariidae species exhibit heterogeneous trends within this broader pattern, with some populations stable or increasing due to factors such as access to anthropogenic food subsidies, while others show sharp declines linked to density-dependent regulation and environmental variability.[73] For instance, the white-chinned petrel (Procellaria aequinoctialis) experienced a reduction from approximately 1.43 million breeding pairs in the 1980s to 1.2 million by 2011, reflecting ongoing pressures amid variable colony-specific dynamics.[74] Breeding success in Procellariidae typically ranges from 60% to 80% in unperturbed colonies, with adult annual survival rates often exceeding 90%, supporting slow but resilient population growth under favorable conditions (λ ≈ 1.02–1.16 across reviewed petrel populations).[73][75] Specific examples include the spectacled petrel (Procellaria conspicillata), whose breeding population has increased post-2000 monitoring, leading to an IUCN downlisting from Critically Endangered to Vulnerable in 2007, and the black petrel (Procellaria parkinsoni), with estimated annual growth rates between -2.3% and +2.5% derived from census data.[76][77] These metrics highlight stochastic influences, such as weather-driven breeding failures, alongside intrinsic limits like deferred maturity (often 5–10 years), which buffer against but do not preclude declines in vulnerable taxa.[78] Over 50% of Procellariidae species are classified as threatened on the IUCN Red List, with trends varying by habitat fidelity; island-nesting species often show steeper drops until predator removal enables recovery, as evidenced by post-eradication increases of 22–23% annually in burrow-nesting petrels on rat-cleared islands.[79][80] Long-term monitoring underscores that while global indices suggest contraction, localized recoveries demonstrate density-dependent rebound potential, independent of uniform anthropogenic forcing, with diverse assemblages re-establishing on restored islands over decades.[79][81]

Assessment of Natural Versus Anthropogenic Threats

Invasive predators, such as rats (Rattus spp.), cats (Felis catus), and house mice (Mus musculus), introduced to breeding islands by human activity, inflict severe demographic impacts on Procellariidae populations, primarily through predation on eggs and chicks. Empirical studies document nest failure rates of 60-90% in affected colonies, with rats alone causing up to 80% chick mortality in species like the Pterodroma petrels.[82] Eradication of these invasives has led to significant recovery in seabird densities, including Procellariiformes, with burrow occupancy increasing by over 50% within years post-removal, confirming their causal primacy over other stressors.[79] In contrast, native predation by species like skuas (Stercorarius spp.) and owls imposes episodic losses but aligns with historical population resilience, as evidenced by stable dynamics in predator-present but invasive-free systems.[83] Fisheries bycatch constitutes another dominant anthropogenic threat, with Procellariidae comprising a substantial portion of the estimated 100,000-300,000 seabirds killed annually in longline fisheries alone, though species-specific rates vary widely due to foraging overlap.[84] Trawl and gillnet interactions add further mortality, potentially numbering in the hundreds of thousands globally, disproportionately affecting shearwaters and smaller petrels.[85] Natural perturbations, including storms and volcanism, have historically driven localized extinctions, such as prehistoric petrel declines in the Macaronesian islands amid eruptive activity, yet these events permitted recolonization and did not precipitate family-wide collapses.[86] Plastic ingestion affects many Procellariidae, with occurrence rates exceeding 50% in examined individuals of species like fulmars (Fulmarus glacialis) and shearwaters, though mass burdens remain low (<5% stomach content) and direct mortality links are infrequent.[87] Light pollution disorients fledglings, increasing grounding rates, while overfishing's prey depletion effects are debated, as discards can subsidize foraging and some populations thrive amid fisheries.[73] Global assessments identify invasives and bycatch as affecting over 100 seabird species each, far outranking climate change or severe weather, which primary-threat only 11% of cases; some analyses emphasize natural oceanographic variability in driving short-term fluctuations over anthropogenic warming signals.[88][89]

Human Interactions and Conservation

Historical Exploitation and Utilization

Rakiura Māori have long harvested sooty shearwater (Puffinus griseus) chicks, termed tītī, primarily for food, employing selective methods that targeted accessible burrows and avoided excessive removal to preserve breeding stocks, a practice rooted in pre-colonial traditions and evidenced as sustainable at pre-industrial intensities.[90] Harvest diaries maintained by some families since the 1950s document annual takes ranging from thousands to tens of thousands of chicks per island, reflecting localized controls that correlated with stable or recovering populations under customary management.[91] Similar traditional exploitation occurred among other indigenous groups, such as hunter-gatherers in southern Chile, where archaeological middens reveal consumption of shearwaters alongside albatrosses and cormorants dating back millennia, though without evidence of widespread depletion prior to European contact.[92] In the North Atlantic, northern fulmars (Fulmarus glacialis) faced intensive historical hunting for meat, eggs, and stomach oil on remote islands like St. Kilda and Grimsey, where 19th-century records describe annual collections exceeding 10,000 birds and eggs by island communities reliant on seabird resources for sustenance and fuel.[93] This exploitation, peaking in the 18th–19th centuries, involved fowling techniques such as cliff netting and egging during breeding seasons, contributing to localized population bottlenecks observable in subfossil records of reduced breeding colonies post-hunting eras.[73] Fulmar oil, extracted from stomach reserves, served as a lamp fuel and medicinal agent, with harvest logs from Iceland and the Faroe Islands indicating yields sufficient to support household needs but occasionally leading to temporary breeding site abandonment when overexploited.[93] Procellariidae species have also been harvested for fisheries bait, particularly shearwaters whose carcasses attracted target fish species, a utilization documented in 19th–20th-century North Atlantic and Pacific logs where birds were culled en masse during migrations to supply longline operations.[94] Such practices, combined with food and oil collection, inflicted empirical declines on insular populations, as evidenced by historical accounts of extirpations on predator-free islands following unchecked egging and chick scavenging, with recovery tied to harvest cessation rather than external factors.[73]

Conservation Interventions and Empirical Outcomes

Eradication of invasive mammals from breeding islands has proven effective for restoring Procellariidae populations, with over 800 successful projects globally enabling seabird recovery.[95] On Marion Island, removal of feral cats in 1991 led to higher breeding success in great-winged petrels (Pterodroma macroptera) and blue petrels (Halobaena caerulea), shifting from suppressed rates under predation to levels supporting population growth.[96] Translocation of near-fledglings to predator-free sites has achieved establishment rates approaching 80% visitation and 76% breeding initiation across seabird restoration events, including Procellariidae species, often within 2-5 years post-intervention.[97] For instance, translocations of Newell's shearwaters (Puffinus newelli) and Hawaiian petrels (Pterodroma sandwichensis) to secure colonies have demonstrated sustained breeding without elevated stress or developmental impacts.[98] The Agreement on the Conservation of Albatrosses and Petrels (ACAP), established in 2001, has driven bycatch reductions through technologies like weighted branch lines, bird-scaring devices, and night setting, with best practices endorsed in 2025 updates showing combined measures as most effective for longline fisheries.[99] These interventions have correlated with population rebounds in monitored Procellariidae, such as record breeding numbers of wedge-tailed shearwaters (Ardenna pacifica) within predator-proof enclosures in Hawaii.[100] Empirical data indicate burrow-nesting procellariids can achieve successful conservation in previously invaded ecosystems via such targeted actions, with breeding success post-predator control reaching 50% at protected sites versus 32% under ongoing predation pressure.[68] Conservation debates highlight the superior cost-benefit of invasive eradication and habitat restoration over expansive marine reserves or climate adaptation measures, given evidence of inherent population resilience in Procellariidae when direct threats like predation and bycatch are addressed.[73] Recent assessments underscore that invasive species exert the most verifiable causal impacts, outperforming speculative climate attributions in driving declines, with post-intervention outcomes like 76% breeding success far exceeding untargeted efforts.[97] A 2024 review of seabird threats prioritizes invasive alien species eradication alongside bycatch mitigation, advocating these over broader environmental interventions lacking comparable empirical validation.[101] Future directions emphasize integrating predator control with translocations, informed by long-term monitoring to maximize recovery while avoiding resource diversion to less substantiated risks.[97]

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