Small seabirds with prominent upturned nostrils and dancing flight over the waves
5.5–10 in (13–26 cm)
Number of genera, species
8 genera; 21 species
Several species, whose status is unknown, are possibly rare
Evolution and systematics
The fossil record is poor, represented only by Oceanodroma hubbsi from the Upper Miocene of California, Oceanites zalascarthmus from the South African Pliocene, and Primodroma bournei from the English Eocene.
A number of studies based on comparative anatomy and DNA analyses suggest that the order evolved close to the base of procellariiform radiation.
The family Hydrobatidae is divided into two subfamilies: the Oceanitinae contains seven very long-legged species found in southern seas, and the Hydrobatinae contains 14 rather shorter-legged species mainly found in northern waters.
Storm-petrels are small, delicate seabirds whose long legs are used to fend off from the water as the birds snap up food items. The wings are rounded at their tips because the tenth and outermost functional primary is shorter than that of the ninth (primaries are main flight feathers). The fused tubular nostrils are prominent and span nearly half the length of the bill. The smallest species, least storm-petrels (Oceanodroma microsoma), weigh only 0.7 oz (20 g) and have a wing span of about 12.6 in (32 cm); the largest, Tristram's storm-petrels (Oceanodroma tristrami), weigh about 2.9 oz (83 g) and have a wingspan of 22.4 in (57 cm).
Storm-petrels tend to have dark blackish or brownish plumage, but many are paler ventrally and have white rumps. The tail may be square cut or forked. The feet of birds of the genera Fregetta and Nesofregetta bear strange spade-like claws. Female storm-petrels tend to be larger than males. Most (perhaps all) storm-petrels carry the musty body odor characteristic of tubenoses. The olfactory lobes of the brain are large and a functional sense of smell has been demonstrated in field experiments at sea and over land.
Storm-petrels are found worldwide, but they are particularly numerous in the vast Southern Ocean. Many breed around Australasia but five species are concentrated around islands from Mexico to California. At sea they occur in all oceans but fail to penetrate Arctic seas.
Marine distributions of storm-petrels are poorly known; being small, storm-petrels are hard to see and identify as they dart along hugging the waves. Some species prefer warm or cool waters. For example, Leach's storm-petrels (Oceanodroma leucorhoa) inhabit cooler water well offshore of the western United States, and wedge-rumped storm-petrels (Ocean-odroma tethys) and Elliot's storm-petrels (Oceanites gracilis) seem confined to cool waters of the Humboldt Current. Some species congregate along areas of upwellings, as does the band-rumped storm-petrel, (Oceanodroma castro), which prefers warm water and aggregates off Florida and South Carolina along Gulf Stream eddies. Storm-petrels breed on islands that are free of mammalian predators.
Most nests are in burrows, and the nest sites are retained from season to season and form the focus for the maintenance of the same pair bonds from year to year. Most visit their nests only after dark and appear to have little by way of display except for mutual preening. Aerial flight displays and chasing in species like the wedge-rumped storm-petrel and Wilson's storm-petrel (Oceanites oceanicus) have been described. The spectacular performances of the first species take place by day, and this species is evidently adapted to nighttime feeding.
In all species studied the sexes call differently. A variety of churring or whirring sounds is heard from burrows of northern species such as the European storm-petrel (Hydrobates pelagicus). On the Norwegian island of Rost, European storm-petrels flutter around the exhaust of the lighthouse engine, and the birds' calls match the rhythm of the engine. Calls may vary from bird to bird, which suggests a role in individual recognition—important for communication after dark. The voices of the southern Oceanitinae tend to be a higher pitch than those of the Hydrobatinae, and their whistles may have a ventriloqual quality.
Most storm-petrels tend to be solitary at sea but flocking also occurs. Some species, like Wilson's storm-petrels, are highly migratory. White-faced storm-petrels (Pelagodroma marina) from Western Australia migrate north to winter along the convergences of the northern Indian Ocean. Other species, such as Leach's and European storm-petrels, shift south after breeding, the former to the tropical Atlantic and Pacific oceans and the latter mainly to the Benguela Current off Africa. Other North Pacific species also shift south; for example, Matsudaira's and Swinhoe's storm-petrels (Ocean-odroma matsudairae and O. monorhis) from Japanese seas reach the Indian Ocean via the Straits of Malacca and the Timor Sea. Most storm-petrels breeding off California just disperse into local seas, although the black storm-petrel (Oceanodroma melania) moves south to the Humbolt current.
Feeding ecology and diet
Crustaceans are important foods for storm-petrels. For example, euphausids are the most frequent prey item for Wilson's storm-petrels nesting around Antarctica; the euphausid species and its relative importance varies with locale and season. Farther north around the Crozet Islands, the same bird still eats crustaceans, but copepods and cirripedes are relatively more abundant in the diet. The gray-backed storm-petrel (Garrodia nereis) seems to specialize on barnacles that evidently are picked off floating rafts of seaweed.
Storm-petrels have a penchant for oily foods. They snip up oil droplets from the sea but seem to avoid man-made oil slicks, perhaps by using their sense of smell. Stomachs usually contain the stomach oil found in most tubenoses, and this oil, being digestible and full of energy, forms an important food for adults and chicks. It is derived directly from prey items, many of which contain heavy loads of oil droplets (especially when breeding).
Storm-petrels usually feed solitarily but will congregate around a suitable food source such as a dead seal or squid. Some species associate with pods of whales, and Wilson's storm-petrels ingest whale feces. Ship following is common; the birds eat small prey churned up by propellers. Wake followers tend to attract others, and up to 50 black storm-petrels have been seen combing the wake at one time.
Storm-petrels feed from the top few inches of the sea. They seem to have the ability to stay within bill range of a heaving sea. This ability is aided by their low wing loadings, which means that as the wave heaves, the air above moves with it and so does the bird. Both Wilson's and black storm-petrels will dive to retrieve food. The precise mode of feeding varies with the species, the length of the legs, and other factors. Some species, like Wilson's storm-petrels, hold their wings high while fending off from the surface with both feet, whereas black-bellied storm-petrels (Fregetta tropica) hold their wings out and skip from side to side. Feeding birds face the wind, and if a gale shifts abruptly through 90 degrees to blow straight down the wave furrows, the tiny birds may be unable to feed and be forced far down wind.
Typically monogamous, storm-petrels first visit their natal colonies as prebreeders, and they breed at 4–5 years old. Nesting occurs when surrounding seas provide plenty of food; that is in the spring or summer in middle and high latitudes. Tropical breeders often have an extended laying season.
With few exceptions (e.g., the wedge-rumped storm-petrel in the Galápagos), all activity on land occurs after dark, and for many species the behaviors culminating in mating are unclear. European storm-petrels often crash into one other, apparently deliberately, and then fall to the ground before disentangling themselves. In Alaska, fork-tailed storm-petrels (Oceanodroma furcata) circling overhead call in response to cries from burrows below, suggesting pair bonding. High-speed zigzagging chases have been seen among other species, but, in general, the significance of these maneuvers is obscure. Not much more is known of their behavior on the ground, but using night vision equipment, copulation of fork-tailed storm-petrels has been seen in the nest chamber and on the ground outside. There was little precopulation ceremony other than mutual preening.
Some female storm-petrels feed at sea while producing the single egg; this trek is called the prelaying exodus. In a study of Wilson's storm-petrels, the females of 31 pairs stayed away for 16–18 days, leaving their partners to visit the nests from time to time, perhaps to keep them clear of snow. In other species there seems to be no clear exodus. In Leach's storm-petrels, semen is stored in special glands in the vaginal folds of the cloaca. This arrangement allows fertilization to be delayed while the bird travels to the best feeding area while producing her egg and returning to her nest.
The single egg is ovoid, whitish, and often sprinkled with pinkish dots. It is laid within 24 hours of the female's return and is very large compared to her body size. That of the least
storm-petrel is 29% of her body weight—probably the heaviest egg relative to body size of any bird. Highly migratory species lay over a relatively short period, whereas sedentary birds like the ashy storm-petrel (Oceanodroma homochroa) breeding off California lay over a period of about 100 days.
The nest is a chamber at the end of a tunnel bored into soft ground or in a crevice among rocks. Males seem to be the tunnel diggers and the chambers are often sparsely lined with bits of local vegetation. Gray-backed storm-petrels and white-bellied storm-petrels (Fregetta grallaria) burrow into the fibrous debris at the bases of tussock clumps and so are, in effect, above ground. The microclimate in a burrow is milder than the climate outside, being more humid and warmer in cold climates and cooler than ambient temperatures in tropical regions. The egg is tucked snugly into a bare, well-vascularized incubation patch. Incubation shifts vary from two to three days in cold water species but average 4.5 days in warm water species like the white-faced storm-petrel.
The female usually leaves for the sea within 24 hours of laying. The male takes the first incubation shift and the sexes take turns thereafter. The total time between laying and hatching in continuously covered eggs varies from 38 to 42 days. When eggs are temporarily abandoned, this interval is longer. Eggs are resistant to chilling, can be abandoned for several days, and yet will still produce a chick. In one extreme case, a fork-tailed storm-petrel egg left uncovered for 31 days still hatched. The embryo's resistance to chilling while still remaining viable is a valuable adaptation for a small seabird having to cope with variable weather and sea conditions which may prevent parent's return.
The chick is hatched covered in down, its eyes closed, and with incomplete control of its body temperature. It is brooded by one or other of the parents for three or more days—known as the guard stage. In their thick down the chicks look like powder puffs. They are grayish or whitish, often paler on the belly. In most, the initial down (protoptile) is replaced by a second (mesoptile) down that grows attached to the protoptile, which, in turn, is pushed out by the feathers proper (the teleoptiles). At least two of the southern birds, Wilson's storm-petrels and white-faced storm-petrels, have only one down.
Initially the chick gets small meals, perhaps mostly of stomach oil, but meal sizes increase when the guard stage is over. With the milder climate in the burrow and a layer of subdermal fat, the chick can then thermoregulate with the milder climate in the burrow. Both parents feed the chick and a fairly even division of labor seems to be the norm. Some meals can be huge, e.g., as much as the chick's own body weight (usually after both adults have fed it on the same night). Parental feeding visits are frequent at first but decline in older chicks. For instance, European storm-petrel chicks 11–20 days old received visits on 93% of the nights, those aged 31–40 days on 85% of the nights, and 51–60 day-olds on 66% of nights. Chicks can withstand fasting for 6–7 days. Starved chicks may become torpid to reduce energy needs, but their body weight falls and death may result.
A chick's growth follows a typical logistical curve, climbing steeply for about the first half of the nestling stage then stabilizing above adult weight and finally falling in the last 10–20 days before first flight. This is preceded during its last week or so by its leaving the burrow to exercise its wings and to explore the vicinity of the nest. A chick's first flight usually takes place from some nearby eminence. Parents take no part in this; the chick leaves alone and, as far as is known, once at sea is really on its own. Average nestling periods (the days between hatching and the first flight) range from 57 to 84 days. Nestling periods are much more variable than incubation periods, partly due to erratic provisioning. Breeding success (percent of eggs laid that produce flying young) varies from year to year.
Some species are abundant with many colonies scattered widely. These include Leach's, European, Wilson's, gray-backed, white-faced, and band-rumped storm-petrels. There are also a number of very localized species such as Matsudaira's storm-petrel, which breeds only at the Volcano Islands, and the ashy, least, and black storm-petrels, which nest off southern California and Mexico.
For some species, breeding places are virtually unknown. For example, Markham's storm-petrel (Oceanodroma markhami) has been found nesting on the Paracas Peninsula, Peru, but more undiscovered colonies must exist to account for the numbers of birds encountered at sea. None of these seems to be endangered, but Elliot's storm-petrel may well be as only one pair of the subspecies Oceanites gracilis gracilis has been found nesting off Chile. Breeding places of O. g. galapagoensis remain undetected.
Colonies of storm-petrels have been wiped out by predators, which are usually mammals like rats and mustelids (even mice can damage these small birds by eating their eggs). The extinction of the Guadalupe storm-petrel (Oceanodroma macrodactyla) has been attributed to a combination of predation by cats and erosion from grazing by goats. A particularly noteworthy conservation effort is the extermination of foxes from colonies of Leach's and forked-tailed storm-petrels in Alaska, which resulted in the reestablishment of many colonies. Another success story concerns the use of tape playbacks of Leach's storm-petrel calls plus the provision of artificial burrows on islands in Muscongus Bay, Maine. Attracted by the amplified calls, birds investigated the burrows and eventually bred there. A colony was established (on Ross Island) where there had been no previous evidence of breeding.
No storm-petrel species is threatened, although birds like Hornby's (Oceanodroma hornbyi) and Markham's, being quite unknown, badly need investigation. The populations of Tristram's storm-petrel on Midway Island, formerly decimated by rats, seem to be recovering since the rats were wiped out.
Significance to humans
Storm-petrels were well known to early sailors and very familiar to sealers and whalers because their hunting activities attracted birds like the cosmopolitan Wilson's storm-petrel. They lumped several species together as Mother Carey's Chickens, Mater cara referring to the Virgin Mary under whose protection seafarers were supposed to come. Although killing of these birds was sometimes considered fraught with danger, they were often caught for use as bait. Wilson's storm-petrels were killed at night from the stern of the ship, and many are attracted to lights. Sealers threaded wicks through the alimentary tracts of adults or fat chicks to draw out the stomach oil, which could be used as a candle.
Small though they are, several species have been eaten regularly by primitive societies: Aleuts, American Indians, and the inhabitants of the Izu Islands, Japan, all ate Leach's storm-petrels, while the Morioris of the Chatham Islands, New Zealand and aborigines of eastern Australia ate white-faced storm-petrels when available.
List of SpeciesWilson's storm-petrel
Oceanites oceanicus Kuhl, 1840, no locality. O. o. oceanicus: Islands off Tierra del Fuego and subantarctic islands of Atlantic and Indian Oceans, including South Georgia; O. o. exasperatus: South Shetland, South Sandwich, South Orkney Islands, Elephant Island, coasts and offshore islands of Antarctica.
other common names
English: Mother Carey's chicken; French: Pétrel océanite; German: Buntfüssige Sturmschwalbe; Spanish: Paiño de Wilson.
7 in (18 cm); 1.3 oz (35 g). Wholly black above and below except for white rump merging into the white lower flanks and thighs and a pale band across the center of each wing. Tail square cut and black. Legs black, very long, and projecting beyond the tail when flying; webs between toes yellow. Juvenile like adult. Bill black with prominent nasal tubes reaching about halfway along the ridge of the bill. Sexes alike.
Wholly marine except when nesting, found in all oceans particularly along coastal upwellings and fronts. It tends to be more often seen offshore compared to other storm-petrels such as Leach's and whitefaced storm-petrels, which prefer deeper water.
Highly migratory, moving from April to June from the southern breeding stations to northern reaches of the oceans, but avoids Arctic seas. In the Atlantic the journey from the south is 7,000 miles (11,000 km) for some birds.
These birds are concentrated along the ocean shelves during the northern summer. Although most move back to southern waters to breed during the northern winter, some remain: these are probably juveniles or birds that are taking a season off-duty—a so-called sabbatical year.
The feeding behavior is distinctive with a flight characterized by alternate glides and wing flutterings while the long legs are drooped and often break the surface. Most food is snipped from the surface without alighting, and this is the most common ship's follower. Calls used on the breeding grounds include a grating sound used by both sexes and a chatter call used by the males to attract females.
feeding ecology and diet
Crustaceans, but fish are also eaten (these are more energy rich than crustacea) with mycophids up to 3.3 in (8.5 cm) long being fed to the chicks (quite a meal for a bird with a bill only 0.5 in [1.2 cm] long).
The pair-bond is held over several seasons and most pairs tend to breed annually. The nest forms their focus. There is little to suggest that they stay together during migration. Because of the short polar summers, Wilson's storm-petrels breeding around Antarctica have accelerated the development of the egg and chick: the time from laying to fledging is 91 days (the shortest period for any tubenose). Birds farther north, though slightly smaller, take longer, perhaps because the food supply is less concentrated than it is off the southern continent.
Most nests are hidden in crevices among rocks or coarse scree. The egg is laid on the bare earth in a shallow scrape, and those on the southern islands are often lined with scraps of local vegetation.
The eggs take about 40 days to hatch if continually incubated. The chick flies at 48–78 days old. A major cause of mortality is unseasonal weather that stops birds entering their nests or freezes chicks within them. Predation by skuas is not usually significant.
Not threatened. One of the most abundant seabirds. Its isolation is its major safeguard.
significance to humans
Oceanodroma leucorhoa Vieillot, 1818, maritime parts of Picardy. O. l. leucorhoa: all North Atlantic and North Pacific populations south to California. Populations in eastern Pacific vary in size and degree of white on rump; O. l. chapmani: breeds on San Benitos and Los Coronados islands, Mexico. O. l. socorrensis: breeds in summer on islets off Guadalupe Island, Mexico. O. l. cheimomnestes: breeds in winter on islets off Guadalupe Island, Mexico.
other common names
French: Pétrel culblanc; German: Wellenlaüfer; Spanish: Paiño de Leach.
7.0–8.5 in (180–220 mm); 1.3–1.9 oz (38–54 g). Medium-sized storm-petrel with white rump patch and distinctly forked tail, blackish brown above and below with paler diagonal upperwing bar from carpal joint back to trailing edge near body. Strongly downhooked bill and prominent nostrils. Legs and feet black. Sexes alike. Flight less fluttering than that of Wilson's storm-petrel and wings tend to be held more horizontally than those of Wilson's storm-petrel (which raises its wings into a V). Feet not visible beyond tail and much less prone to pattering.
Breeds on islands in the North Atlantic and North Pacific that lack mammalian predators. Found at sea throughout these seas, migrates south into the tropics after breeding. Pairs have even been found in burrows on the Chatham Islands, New Zealand. One bird was found on St. Croix Island, South Africa, which suggests the possibility of extending the breeding range southwards. The eastern North American populations shift south to Brazilian waters but many cross to European seas like the Bay of Biscay. British breeders appear to winter mainly off tropical Africa. Japanese and Alaskan birds also winter in tropical seas.
Ranges widely in the open sea. California birds feed farther out in warmer, less productive seas than do ashy and fork-tailed storm-petrels with which they often share nesting islands.
On land, overflying birds that emit calls are mainly prebreeders; nesting birds call mostly from their burrows. Little display occurs between breeding birds except persistent calling using two main types of rhythmic purrings and chatterings. Chatter calls are given from the air and the burrow, and research in Japan suggests that variation in the pitch of calls among birds of the same sex may be used for individual recognition.
feeding ecology and diet
Nekton and planktonic organisms are taken from the surface while the bird hovers facing the wind, sometimes alighting momentarily. Otherwise the flight is a mixture of gliding and rather wild dashes, with many changes of direction. Seldom follows ships but is prone to be wrecked on beaches during gales.
Diet includes a great range of fish, crustaceans, and squid as available. The birds tend to seek areas where upcurrents bring organisms to the surface. Their stomachs often contain deep-sea animals that only approach the surface at night, evidently taken after dark. They appear to find some prey using their good sense of smell.
Most breed at around five years old. Having gained a nest site, a pair remains intact as long as they reproduce satisfactorily. Most dig burrows, but some occupy crevices among rocks or stone walls. The single egg is occasionally replaced if it is lost soon after laying. Both sexes incubate in 2–3 day shifts for about 43 days. Chick is fed almost nightly in the first few days after the brief brooding period, then less often as it grows. Unfed chicks become torpid but can recover. Growth follows the normal curve: weight climbs steadily at a constant rate, levels out at above parental weight, then falls in the last 10 or so days before fledging.
Fledging occurs when the chick is between 56 and 79 days. The number of chicks fledged per egg laid differs between seasons and places and ranges from 48 to 73%. Losses have often been due to introduced mammals like mink, cats, and others, but natural predators also include owls, eagles, corvids, and gulls.
Not threatened, but existing colonies need protection against the introduction of placental mammals and trampling of burrows.
significance to humans
Ainley, D.G., and R.J. Boekelheide, eds. Seabirds of the Farallon Islands. Stanford, CA: Stanford University Press, 1990.
Huntington, C.E., R.G. Butler, and R.A. Mauck. "Leach's Storm-petrel." In The Birds of North America, No. 223, edited by A. Poole and F. Gill. Philadelphia: Academy of Natural Sciences; Washington, DC: American Ornithologists' Union, 1996.
Lockley, R.M. Flight of the Storm-petrel. London: David and Charles, 1983.
Warham, J. The Behaviour, Population Biology and Physiology of the Petrels. San Diego: Academic Press, 1996.
Warham, J. The Petrels: Their Ecology and Breeding Systems. San Diego: Academic Press, 1990.
Croxall, J.P., H.J. Hill, R. Lidstone-Scott, M.J. O'Connell, and P.A. Prince. "Food and Feeding Ecology of Wilson's Storm-petrel Oceanites oceanicus at South Georgia." Journal of Zoology, London 216 (1988): 83–102.
Taoka, M., T. Sato, T. Kamada, and H. Okumura. "Sexual Dimorphism of Chatter-Calls and Vocal Sex Recognition in Leach's Storm-Petrels (Oceanodroma leucorhoa)." Auk 106(1989): 498–501.
Warham, J. A Bibliography of the Procellariiformes or Petrels. February 1999. (January 31 2001). <http://www.zool.canterbury.ac.nz/jwjw.htm>
John Warham, DSc