Gaviiformes

Loons

(Gaviidae)

Class Aves

Order Gaviiformes

Family Gaviidae

Number of families 1

Thumbnail description
Medium to large-sized, foot-propelled diving waterbirds that feed mainly on fish. Foot placement is far posterior, making walking on land difficult. Bills are sharp and dagger-like. Alternate (breeding) plumage is boldly patterned primarily with black, white, and gray; nonbreeding plumages are drab gray-brown and white. All have brilliant red iris in alternate plumage, and distinctive eerie vocalizations

Size
20.8–35.8 in (53–91 cm); 2.2–14.1 lb (1.0–6.4 kg); males slightly larger than females in all species

Number of genera, species
1 genus; 5 species

Habitat
Breed in forested and tundra lakes and ponds, winter at sea and large reservoirs

Conservation status
No species considered Endangered or Threatened

Distribution
Holarctic

Evolution and systematics

The five extant species currently recognized are descendants of an ancient bird lineage. The extinct genus Colymboides arose in the late Eocene to early Miocene. Gavia appeared in the Miocene, and radiated into three size classes by the early Pliocene. Biochemical analyses suggest that loons are most closely related to penguins (Sphenisciformes), tubenoses (Procellariiformes), frigatebirds (Fregatidae), and possibly auks and gulls (Charadriiformes). Traditionally loons have been grouped with grebes (Podicipediformes) because the two orders are convergent. Within the family, common (Gavia immer) and yellow-billed (G. adamsii) loons are very closely related, as are Arctic (G. arctica) and Pacific (G. pacifica) loons. Red-throated loons (G. stellata) are considered less closely related to the other four species, but phylogenetic relationships are uncertain and controversial.

Physical characteristics

Loons are medium to large, foot-propelled diving waterbirds, with anatomy specialized for pursuit and capture of fish. Overall shape is very distinctive with a short neck, pointed wings, and legs set far back on the body. Loon tarsi are flattened and knife-like, cutting through the water efficiently. The feet are large and palmate with the front three toes webbed, and a free hallux. Loon feet, legs, and nails are uniquely countershaded such that when swimming white surfaces, the tops of their feet are oriented down, helping the bird blend in against light sky. Bills are medium-sized and dagger-like. All species have a distinct blood red iris in alternate plumage. Sexes are similar, with the male larger than the female in all species. Each species has four plumages after they are fully grown: juvenal, second alternate (second summer), basic (nonbreeding), and alternate (breeding). Molt times are slightly different for all species.

In alternate plumage all species have striking patterns composed of black, white, and gray. Upperparts are dark gray or black, with faint white speckling to bold white checkering. Underparts are completely white. Each species has a series of bold, thin, black-and-white parallel stripes on the neck. Head patterns are similar in Arctic/Pacific loons and common/yellow-billed loons. The head pattern of red-throated loon is unique: it is the only loon to have brick red in its plumage.

All loons have similar juvenal, second alternate, and basic plumages. Upperparts are gray-brown and underparts are

white as a rule; head pattern is slightly different for all species. Juvenal and second alternate birds appear more scaly than adults due to the presence of pale fringes on many contour feathers. Identification of non-breeding birds can be difficult, and general body shape, bill shape and size, and posture can be more helpful than plumage differences for identification.

Molt is complicated and not yet fully understood. Loons have one complete pre-basic molt in fall and one partial pre-alternate molt in spring. The complete molt in fall is prolonged, and individuals may appear to be molting much of the year. Due to the high wing loading in loons, flight is impossible with the loss of a few primaries. Instead of a gradual molt, which would leave them flightless for months, loons have evolved to molt all of their primaries simultaneously, which only leaves them flightless for a couple of weeks. This molt occurs during winter, from about January to April in all species but the red-throated loon, which is small enough that it can fly during a gradual molt of flight feathers. First-year birds also molt primaries simultaneously in their first summer on salt water.

Distribution

Holarctic in distribution, loons breed from north temperate areas to the high arctic. All species migrate up and down coasts and across land to winter primarily at sea, south to coastal Baja California, Gulf of Mexico, Mediterranean Sea, and coastal China. Loons may stage on inland lakes and rivers during migration. Nonbreeding birds will often spend summer in their species' winter range.

Habitat

Loons breed in freshwater inland lakes and tundra ponds. Where sympatric, different species occupy different-sized lakes. Larger species exclude smaller species from breeding ponds, but are limited to larger ponds by minimum take-off distances. Smaller species can occupy ponds too small for larger species. Individuals usually spend winter near shore in oceans and seas, less than 62 mi (100 km) offshore. They are occasionally found in large freshwater lakes and rivers during winter.

Behavior

Loons are extremely territorial on their breeding grounds. Pairs have been observed attacking their own species, as well as other species of loons, ducks, and geese. In one study, 50% of common loons had healed fractures believed to have been caused by the bill of other common loons. Loon pairs have a series of territorial threat postures and calls to prevent fighting, which can be fatal. In winter and during migration loons may be found singly or in loose flocks.

Loons are known for their unusual vocalizations, described as yodels, wails, and tremolos. Most vocalizations are given on the breeding grounds, occasionally during winter, or on migration. Territorial yodels, most often given at night or early morning, can be heard from great distances (reports have been made of calls heard up to 16 mi [25.8 km] by humans). These male yodels are individually recognizable throughout the birds' lifetime.

Flight is powerful and direct, with wings beating constantly. Most species require running on the surface of water to gain speed for take-off. The red-throated loon is the only species that can take off from land. All species are awkward on land; the posterior positioning of the feet often forces them to push themselves along breast first on their bellies. Occasionally loons accidentally land on wet pavement after mistaking it for water, or are forced to land during storms. In this case, they are stranded and will most likely die, although adults in this situation have traveled over a kilometer to seek water at the expense of broken toes and wrists.

The agility of loons on water compensates for their inadequacies on land. Posterior placement of the feet allows for powerful swimming and diving capabilities. Adults can stay submerged for several minutes and can travel hundreds of meters underwater. Loons dive with a forward thrust using both feet simultaneously to propel them through water. Occasionally, wings are also used to supplement the feet underwater, and they can use one foot as a rudder when turning.

Feeding ecology and diet

Loons take a variety of vertebrate and invertebrate foods, but small to medium-sized fish up to about 7–8 in (18–20 cm) are the primary items. Young are also fed crustaceans, mollusks, and worms. Prey is located by sight from the surface of the water. Loons peer downward, often with their bill in the water, and dive with a thrust from both feet. Most prey is eaten underwater; fish that are too large to be handled underwater are taken to the surface. Most foraging is done close to the surface, but loons may forage as deep as 230 ft (70 m) if the water is clear enough. Very small serrations on the bill help loons hold onto prey. Adults and young consume large quantities of prey; an adult common loon consumes 1,214 kilocalories a day, and a pair may eat 2,000 lb (910 kg) of fish in a breeding season. Loons have large salt glands that remove excess salt consumed from marine environments.

Stomach contents have shown that loons consume a wide variety of fish, including sticklebacks (Gasterosteus), trout (Salvelinus), sculpin (Leptocottus), cods (Gadus), herrings (Clupeiidae), haddock (Melanogrammus), whitefish (Coregonus), capelins (Mallotus), minnows (Cyprinidae), and many other species. At sea, menhaden (Brevoortia) are extremely important. Little is known about prey selection.

Reproductive biology

Loons are monogamous, although they will quickly replace a lost mate. Extra-pair copulation has been noted with marked birds, but has not been studied extensively to determine frequency. Pair bonds, which may last for life, are first formed on breeding grounds. Pair formation is not well understood, although it includes bill-dipping and paired-swimming displays where both adults rise out of the water in various postures. Both sexes quickly build the nest, sometimes in as little as half a day to a week; returning pairs may reuse old nests. Nests are constructed of wet vegetation on land or as a floating mat, with a 15–32 in (38–82 cm) diameter. Two eggs are usually laid (rarely one or three) from May to July, depending on latitude and weather; in northernmost areas there may be only two to three months to breed. Eggs are long, subelliptical in shape, from 2.9 by 1.8 in (72.7 by 44.8 mm) to 3.5 by 2.2 in (89.4 by 55.15 mm) in size, and are olive green with brown markings. Pairs will re-lay if the first clutch is lost. Both sexes incubate, beginning with the laying of the first egg. Incubation period is 24–30 days; larger species have longer incubation periods. Chicks hatch asynchronously, are semi-precocial, and covered with dark gray natal down. Chicks leave the nest soon after they dry, but may return for brooding. Young rely on both parents for food but will begin to dive on their own in three days. Chicks will occasionally ride on a parent's back when small. Young can fly in six to eight weeks. Predators of eggs and chicks include many mammals and other birds. To avoid predators, chicks dive to the bottom of the water to stir up sediments, resurfacing in emergent vegetation for cover. Adults have few predators, and band recoveries suggest loons have a life span of 25–30 years.

Conservation status

No loons are listed on the IUCN Red List of Threatened Species of Birds. Isolation of breeding habitat protects their

numbers from human disturbance in many areas, but where overlap does occur loons have declined due to habitat encroachment and acid precipitation, which can lower pH levels enough to kill all the fish in many lakes. Loon conservation groups have formed to protect them in many areas. At sea, fishnets kill many adults, and are responsible for one-third of red-throated loon banding recoveries. Oil spills kill many loons, which are especially susceptible when adults are flightless while molting primaries. The Exxon Valdez spill in Alaska resulted in hundreds of loons being washed ashore. Botulism, a bacterial disease, has killed thousands of loons staging on the Great Lakes. Common loon populations, in particular, have declined in parts of northeastern United States and in Ontario, Canada, due to increase in human activities—boating, use of jet skis, and canoeing—lake acidification, and mercury poisoning.

Significance to humans

The Inuit legally hunt loons in arctic North America for subsistence purposes. Roughly 4,600 may be taken each year. They are not considered to be the best-tasting food, and may be fed to dogs. Native Americans honor loons with many stories and parables. Loons are a symbol of the north, and a symbol of tranquility. The common loon is featured in Canadian currency on the $1 coin (commonly called "loonies") and the $20 bill.

Species accounts

List of Species

Red-throated loon

Gavia stellata

taxonomy

Colymbus stellatus Pontoppidan, 1763, Tame River, Warwickshire, England. Monotypic.

other common names

English: Red-throated diver; French: Plongeon catmarin; German: Sterntaucher; Spanish: Colimbo Chico.

physical characteristics

20.8–27.19 in (53–69 cm); 2.2–5.9 lb (1.0–2.7 kg). The smallest and least robust in the family, with proportionally smaller, upturned bill and smaller feet than other loons. Smaller size allows red-throated loons to take off directly from water and even from land. In alternate plumage, has grayish upperparts, white underparts, gray face, and brick red throat patch. In basic plumage, has grayish upperparts with white speckling, gray cap and nape, white underparts, throat, and face. Juvenal and second alternate plumages similar to basic plumage, with gray-brown wash on head and neck.

distribution

Breeding range is circumpolar, ranging farther north than other loons. Occupies coastal plain in Alaska, northern Canada, Greenland, Iceland, northern British Isles, Norway, Sweden, Finland, and across Russia. Winters on coasts on Atlantic and Pacific Oceans north of the tropic of Cancer, occasionally found inland. Migrates mostly along coast, occasionally over land.

habitat

Breeds mainly on ponds in coastal tundra, occasionally inland up to 3,511 ft (1,070 m) in elevation. Where it competes with other loons, occupies smaller (sometimes fishless) ponds too small for larger loons. In the far north where it is the only loon present, will breed on larger ponds and lakes. Winters on coasts, usually within 3 mi (5 km) of shore in areas with a soft, sandy substrate. Occasionally found inland on large lakes and rivers.

behavior

The only loon to have duet vocalizations, given by pairs on breeding ponds. May migrate singly or in loose flocks. Does not require running start from water during take-off like other loons, and is the only loon that can take off from land.

feeding ecology and diet

Feeds on variety of small freshwater and marine fish. Will feed invertebrates to small chicks, and will feed on invertebrates as adults when fish are scarce. When breeding in fishless ponds, will fly to the coast and other ponds to catch prey to bring back to the young.

reproductive biology

Breeds from May to September, depending on latitude and climate. Incubation 24–27 days. Occasionally moves from breeding

pond to a larger pond or the ocean. Chicks are more agile on land than are adults, and have been seen traveling over a kilo-meter over land. Young can fly after 38 days. Predators include Arctic fox (Aloplex lagopus) and other mammals, jaegers (Stercorarius), and gulls (Larus).

conservation status

Declining over much of its range, although the cause is unknown. Not listed on IUCN Red List of Threatened Birds.

significance to humans

Inuit legally hunt around 4,600 loons (of all species) each year for subsistence; the proportion that are red-throated is unknown.

Pacific loon

Gavia pacifica

taxonomy

Gavia pacifica Lawrence, 1858. Monotypic.

other common names

English: Pacific diver; French: Plongeon du Pacifique; German: Weissnackentaucher; Spanish: Colimbo del Pacifico.

physical characteristics

20–27 in (50–68 cm); 3.7 lb (1.7 kg). Very similar to the larger Arctic loon in all plumages. Bill medium-sized, straight. Black upperparts with white patches, white underparts, black throat, and gray head and neck (darker near bill). Differentiated from the Arctic loon by black flanks, paler nape, and thinner white stripes on neck. In hand, throat shows faint purple iridescence. Basic, juvenal, and second alternate plumages similar, with gray upperparts, crown, and nape and white underparts.

distribution

Breeds on tundra ponds from eastern Siberia to Hudson Bay; winters on Pacific Ocean from southern Japan to Siberia, Alaska to southern California. Migration chiefly occurs coastally.

habitat

Breeds on medium-sized lakes and ponds in northern forests and tundra. Excluded from large lakes by yellow-billed and common loons; excludes red-throated loons from medium-sized lakes. Winters in coastal areas, often farther offshore than other species.

behavior

Males give territorial yodel call that is individually recognizable, and can be heard from miles away. Pacific loons are unafraid of humans, and allow close approach on breeding grounds. A pair was observed adopting a brood of spectacled eiders; this is the only reported case of adoption in Gaviidae.

feeding ecology and diet

Consumes a wide variety of fish; feeding and diet similar to other species.

reproductive biology

Breeds on medium-sized ponds, sympatric with the Arctic loon where ranges overlap; pairs of both species have been found on the same lake. Breeds from May to September. Incubation 28–30 days, fly at 60 days. Predators include gulls (Larus), foxes (Aloplex and Vulpes), jaegers (Stercorarius), and ravens (Corvus).

conservation status

Most populations stable. Not listed on IUCN Red List of Threatened Birds.

significance to humans

Inuit legally hunt loons for subsistence on breeding grounds; 4,600 loons (of all species) are taken yearly.

Arctic loon

Gavia arctica

taxonomy

Colymbus arcticus Linnaeus, 1758, Sweden. Two subspecies recognized.

other common names

English: Black-throated diver; French: Plongeon Arctique; German: Prachttaucher; Spanish; Colimbo Arctico.

physical characteristics

23.6–29.6 in (60–75 cm); 5.7 lb (2.6 kg). Very similar to the smaller Pacific loon. Black upperparts with white patches, white underparts, black throat, and gray head and neck (darker near bill). Differentiated from the Pacific loon by white flanks, darker nape, more distinct white stripes on neck. In hand, throat shows faint greenish iridescence. Basic, juvenal, and second alternate plumages similar, with gray upperparts, crown, and nape and white underparts.

distribution

Breeds from extreme western Alaska across northern Eurasia to northern Scotland. Winters coastally from Japan to China and Europe.

habitat

Breeds on medium-to-large lakes and ponds in northern forests and tundra; winters on coasts.

behavior

Similar to the Pacific loon.

feeding ecology and diet

Consumes a wide variety of fish, similar to other loons. Has been observed catching frogs.

reproductive biology

Incubation 28–29 days, flight 60 days. In Scotland, artificial breeding platforms have been used to increase chick production by an estimated 44%.

conservation status

Most populations stable, not listed on IUCN Red List of Threatened Birds.

significance to humans

A species of interest for many people. Nest platform programs have been started in Scotland, where populations have declined.

Common loon

Gavia immer

taxonomy

Colymbus immer Brünnich, 1764, Faeroes. Monotypic.

other common names

English: Great northern diver; French: Plongeon huard, Plongeon Imbrin; German: Eistaucher; Spanish: Colimbo grande, Colimbo Comun.

physical characteristics

26.0–35.8 in (66–91cm); 5.5–13.4 lb (2.5–6.1 kg). Very similar to the yellow-billed loon in all plumages. In alternate plumage, black upperparts with white checkering and spotting, black neck with white stripping, and black head. Underparts are white. Juvenal, second alternate, and winter plumages similar, dark gray brown upperparts, head, and nape; white underparts and throat. Bill is straight and black in alternate and dark gray with a black culmen in other plumages.

distribution

Breeds throughout Alaska, Canada, northern New England, northern Midwest, and parts of Greenland and Iceland. Winters in Pacific Ocean from southern Alaska to Baja California, on Atlantic Ocean from Newfoundland to Mexico, also Europe and Iceland. Migrates over land and down coasts; stages on larger lakes. Many non-breeders summer in winter range.

habitat

Breeds in clear, oligotrophic, forested lakes and large tundra ponds. Winters mainly on coast within 62 mi (100 km) of shore, occasionally on large inland lakes and rivers.

behavior

Found in pairs on breeding grounds, singly or in loose flocks during migration and winter. Requires 100–650 ft (30–200 m) to take off, limiting common loons to large lakes. Extremely territorial on breeding grounds—other loons and waterbirds are chased off. Yodel call, a series of repeated two-note phrases, is recognizable to individuals and used to defend territories.

feeding ecology and diet

Feed mainly on fish and invertebrates; vegetation occasionally taken. Crayfish are a common food when fish are scarce.

reproductive biology

Nests farther south than other loons, from May to October. Little is known about pair formation. Nest is made of vegetation in about a week, on land at the edge of a lake. Incubation 27–30 days. Young leave nest in one day, but may return for brooding. Young able to fly in 11 weeks. Predators include gulls (Larus), ravens and crows (Corvus), pike (Esox), and raccoons, weasels, and skunks (Carnivora).

conservation status

Populations are stable. Not listed on IUCN Red List of Threatened Birds, but is listed as threatened or of special concern in several northeast states. Acidification of lakes, heavy metal contamination, and human encroachment threaten populations in southern range.

significance to humans

Inuit hunt 4,600 loons (of all species) per year for subsistence. Many Native American tribes have stories about common

loons. Many loon conservation groups have also been formed to protect common loons in their southern range.

Yellow-billed loon

Gavia adamsii

taxonomy

Colymbus adamsii Gray, 1859, Alaska. Monotypic.

other common names

English: White-billed diver; French: Plongeon a Bec Blanc; German: Gelbschnabel-Eistaucher; Spanish: Colimbo de Adams.

physical characteristics

30–36 in (76–91cm); 9.0–14.1 lb (4.1–6.4 kg). Largest loon, with the largest bill. Very similar to the common loon in all plumages. In alternate plumage, black upperparts, neck, and head; white striping on the neck and white checkering on the back. Underparts white. Juvenal, second alternate, and winter plumages similar, dark gray brown upperparts, head, and nape; white underparts and throat. Bill is large and upturned, yellow to ivory.

distribution

Replaces common loon in high arctic, breeds in Canada and Alaska, also across Siberia. Winters along coasts in northern Pacific, and along the coast of Norway. Winters farther north than other loons.

habitat

Breeds in large tundra lakes; winters along coasts.

behavior

Similar to the common loon.

feeding ecology and diet

Up-turned bill has been suggested as an adaptation for feeding along water bottom; very little data on diet, probably similar to other loons.

reproductive biology

Nests from June to September on hummocks near breeding ponds. Incubation around 28 days; young leave nest within one day of hatching. Young can fly at about 11 weeks. Predators include foxes (Aloplex), ravens (Corvus), and gulls (Larus).

conservation status

Rarest of the loons, but not threatened. Populations stable.

significance to humans

Taken occasionally for subsistence by Inuit.

Resources

Books

Harrison, C. J. O., and P. Castel. Bird Nests, Eggs, and Nestlings of Britain and Europe. Milan: New Interlew Spa, 1998.

Josson, Lars. Birds of Europe with North Africa and the Middle East. Princeton: Princeton University Press, 1993.

Wild Bird Society of Japan. A Field Guide to the Birds of Japan. Tokyo: Wild Bird Society of Japan, 1985.

Periodicals

Abraham, Kenneth F. "Adoption of Spectacled Eider Ducklings by Arctic Loons." Condor 80, no. 3 (1978): 339–340.

Barr, Jack F., Christine Eberl, and Judith W. McIntyre. "Red-Throated Loon (Gavia stellata)." Birds of North America no. 513 (2000): 1–28.

Hancock, Mark. "Artificial Floating Islands for Nesting Black-Throated Divers (Gavia arctica) in Scotland: Construction, Use and Effect on Breeding Success." Bird Study 47, no. 2(2000): 165–175.

McIntyre, Judith W., and Jack F. Barr. "Common Loon (Gavia immer)." Birds of North America no. 313 (1997): 1–32.

North, Michael R. "Yellow-Billed Loon (Gavia Adamsii)." Birds of North America no. 121 (1994): 1–24.

Vallianatos, Mary, and Jon McCracken. "Loons and Type E Botulism." Birdwatch Canada 15 (2001): 9.

Woolfenden, Glen E. "Selection for a Delayed Simultaneous Wing Molt in Loons (Gaviidae)." Wilson Bulletin 79, no. 4(1967): 416–420.

Peter Andrew Hosner