Description
SUMMARY: A large-bodied, elongate loon with a heavy bill. Breeding adults have a black head and bill and white checkering on the back. Juveniles and winter adults are rather plain gray on the upper side and white below, with an irregular edge to the gray-white junction on the side of the neck. Wingspan around 46 inches (117 cm).
DETAILS: Plumage differences between the sexes are indistinguishable. In alternate (breeding) plumage, the head and neck are velvety black with a slight greenish gloss. Across the throat is a prominent transverse bar of short, vertical white streaks; on either side of the neck is a collar of similar, longer white streaks, not quite meeting in the center of the throat or the back of the neck. The bill is black, and the iris is brownish ruby. The upperparts are black with a greenish gloss, heavily spotted with white, each feather (except the unmarked upper tail coverts) with a pair of white, squarish subterminal spots, smallest on the upper mantle, back and rump, and largest on the scapulars. The underparts are mainly white, the sides of the breast are streaked black and white, and the flanks are black with small white spots. The tail is short, with 16-20 feathers, and entirely black. The wings are narrow and pointed, about 35 cm long (unflattened), with 11 primaries and 22-23 secondaries. The remiges are blackish with dark shafts, and the tips of the inner secondaries have single or paired white spots. The coverts are similar to the upperparts but have more circular subterminal spots. The wing lining is mostly white. The legs are black on the outer side paling to grayish or white inside; the webs of the feet are mostly white above, and black with white centers below (Palmer 1962, Jackson 1976, Johnsgard 1987, McIntyre 1988).
Although some overlap exists between the sexes, within pairs, males are consistently larger than females (Barr 1973, McIntyre 1988). Adults measure about 1 m in length outstretched (Palmer 1962, McIntyre 1988). Weight varies geographically and within populations, with adults ranging from 2.7 kg to more than 6.3 kg (McIntyre 1988). Sexual dimorphism is most pronounced in bill length and depth (McIntyre 1975, Storer 1988). Some structures of the digestive tract are larger in males, suggesting possible food partitioning by pair members (Barr 1973).
The definitive alternate plumage is acquired by a complete molt between January and March, which renders birds flightless for nearly a month (Woolfenden 1972, McIntyre 1988). This plumage is not acquired until the third or fourth summer (McIntyre 1986).
The basic (winter) plumage is acquired by a partial molt of contour feathers beginning in late summer and lasting through fall (although earlier in unsuccessful breeders and immatures). In this plumage the forehead, crown and back are grayish-brown, and the chin, throat and foreneck are white. The bill is brownish-gray to pale bluish-gray or horn colored. The iris is brown. The upperparts are brownish-gray, the feathers margined with paler gray, with a few black and white feathers occasionally retained in adults. The underparts are mainly white, with a brownish, streaked appearance on the sides of the breast and flanks. The tail is dark brown, tipped with white (Bent 1919; Johnsgard 1987; McIntyre 1986, 1988).
The juvenal plumage begins to emerge at about one month. This plumage is similar to the adult basic plumage, although the upperparts have paler and more conspicuous feather margins than those of adults, and the throat and sides of the neck are more finely streaked with brown. Flight feather growth is completed at 12-13 weeks of age, and this entire plumage is worn until the following summer when a complete molt produces another, more adult- like basic plumage (Palmer 1962, McIntyre 1988).
Newly-hatched chicks undergo two successive changes of downy plumage. The first down is blackish, paler on the throat, upper breast and flanks, and white on the lower breast and belly. This is replaced at 10-14 days of age by the second downy plumage of primarily brownish-gray feathers, which are replaced by the juvenal contour feathers at four weeks (Palmer 1962, McIntyre 1988).
NESTS: Nests are nearly always built at the water's edge. Substrates range from masses of aquatic and terrestrial vegetation, to moss, to bare soil, sand or rock, to depressions in old muskrat (ONDATRA ZIBETHICUS) houses (McIntyre 1988). Nests are typically large, bulky structures composed of vegetation, if available, but may consist simply of scrapes in the bare soil or duff, or may be placed directly on rocky substrate (Bent 1919, McIntyre 1988). Some are built on sedge or ericaceous bog mats, and occasionally partially rotted, semi-submerged logs are used (McIntyre 1988). Nest materials often include clumps of partially decayed aquatic vegetation, roots and rhizomes (McIntyre 1975, 1988). Nest diameters average 56-66 centimeters outside, 24.5-33 centimeters inside, and 3-7.6 centimeters deep (Olson and Marshall 1952, McIntyre 1975, Sutcliffe 1980).
EGGS: Subelliptical to ovoid in shape and vary from deep-olive to light-brown in color, most being deep-olive brown with irregular dark brown or black spots. Dimensions average 86.5-91.5 mm by 54-57 mm, and weights average 140-160 g.
VOCALIZATIONS: See Barklow (1979), Klein (1985), McIntyre (1988), Miller (1988), Palmer (1962), Rummel and Goetzinger (1975), Sjolander and Agren (1972) for descriptions of vocalizations.
Diagnostic Characteristics
See Stallcup (1994) for information on identification of North American loons.
Habitat
Breeding habitat includes usually clear lakes (McIntyre 1988) containing both shallow and deep water areas (McIntyre 1975, 1988; Strong 1985). In studies comparing lakes with and without loons, higher turbidity was suggested as a factor influencing lack of occupancy (Barr 1973, McIntyre 1988). Nest sites are on small islands(Olson and Marshall 1952, Vermeer 1973, McIntyre 1975, Titus and VanDruff 1981, Strong 1985), quiet backwaters (Strong 1985), or mainland shores. Loons have been found nesting in marshy portions of lakes in water depths no greater than 0.5 m (Alvo 1981). Optimal nest sites, as measured by degree of success, include overhead cover to conceal eggs from predators, protection from wind and waves, good visibility by incubating adults, and a steep slope adjacent to the nest for adequate underwater approaches and exits (McIntyre 1975, 1983, 1988). Brood-rearing areas are typically located in shallow coves of fairly uniform depth, sheltered from prevailing winds and wave action, and are independent of nest site location (McIntyre 1983, Strong 1985). Adults tending chicks prefer shallow water areas (< 2 m) close to land (< 150 m) (Strong 1985, Strong and Bissonette 1989). Deepwater areas (> 4 m) distant from land (> 250 m) are avoided by feeding adults and adults tending chicks, but are often used for social interactions (Strong 1985). Breeding adults usually feed outside of nursery areas (Strong 1985), occasionally outside of their territories (McIntyre 1983), and may visit nearby lakes for feeding (Miller and Dring 1988).
In winter and during migration, common loons use inland lakes and rivers and marine and estuarine coastal waters. Most nonbreeding subadults apparently remain in coastal areas during breeding season. Winter primarily in coastal marine habitats, including bays, coves, channels, inlets and other shallow areas (Bent 1919, McIntyre 1988, Palmer 1962). Some individuals overwinter on inland lakes and rivers, although this appears to be largely weather influenced (McIntyre 1988). While shallow, inshore waters appear to be utilized more frequently than deeper, offshore waters (McIntyre 1978, Daub 1989), some use continental shelf waters up to 100 m in depth and 100 km from land (Haney 1990). In the southeastern U.S. (between 29 degrees and 35 degrees North latitude), wintering loons were most common in waters up to 19 m deep but were rare or absent in highly turbid waters five to 15 km from shore. Loon distribution shifted farther offshore during midwinter to avoid increases in these turbid water areas (Haney 1990). Feeding typically occurs in water depths less than five meters, while maintenance activities (e.g., preening and drifting) take place in deeper water (McIntyre 1978, Daub 1989).
Ecology
Hectares of water area per territorial pair: 503 (New Hampshire); 44, 73 (Minnesota); 39 (Saskatchewan); 351 (New York) (Johnsgard 1987). Lakes smaller than 80 ha generally support only one breeding pair. Typically, territory size is larger on large lakes than on small lakes. Generally, loss of eggs to predators is not a primary cause of breeding failure (Johnsgard 1987). Wintering birds may defend feeding territories during the day, gather into rafts at night.
Ecology of wintering loons is not well studied. McIntyre (1978) found that loons off the Virginia coast maintained individual feeding territories of four to eight ha during the day and rafted together at night. Activity patterns were significantly correlated with tidal changes. Maintenance behavior was greatest during the mid-period of tidal rise. Feeding activities peaked late in the flood tide and during the first half of the ebb tide. In Rhode Island, no winter feeding territories, feeding assemblages, or tide-correlated activity patterns were noted by Daub (1989).
Reproduction
BRIEF SUMMARY: Egg-laying begins one to several weeks after spring arrival, usually during mid-May in the south, and well into June farther north. Replacement clutches may be initiated as late as early July. Incubation lasts around 4 weeks. Chicks leave the nest within 24 hours of hatching and are soon moved to nursery areas. Chicks may be carried on their parents' backs until they reach three weeks of age. Most juveniles are capable of flight at 11-12 weeks, and some leave their small, natal lakes or parental territories shortly afterward.
ARRIVAL AND TERRITORY ESTABLISHMENT: Timing of spring arrival is correlated with latitude and dictated primarily by ice-out phenology (McIntyre 1988). In southern portions of the breeding range, pairs may reoccupy territories in March, while at northern latitudes arrival may be delayed until mid or late May (McIntyre 1988). In Minnesota, an average of eight days elapsed between ice break-up and loon arrival in an early ice-out year, five days in an average year, and three days in a late year (McIntyre 1975). Males typically return first, especially in southern breeding areas (McIntyre 1975, 1988; Sutcliffe 1980). However, pairs often arrive together at northern lakes (McIntyre 1988). Territories are established immediately after arrival and may change in size as the breeding season progresses, expanding after chicks hatch and shrinking for failed pairs (McIntyre 1988).
COURTSHIP: It is believed that pairs remate each spring and that courtship serves primarily to renew the pair bond (McIntyre 1988). Courtship begins shortly after territory reoccupation and involves quiet, shared displays, including simultaneous swimming, head posturing and short dives. Vocalizations are not extensive. Copulation sequences are stereotyped, typically last from three to ten minutes, and take place on land (McIntyre 1988). Some copulation sites become nest sites (McIntyre 1975).
NESTING PERIOD: Nest-building is conducted by both members of the pair and may immediately follow copulation, sometimes lasting over four days (McIntyre 1975, 1988). Egg-laying begins one to 4.5 weeks after spring arrival, usually during mid-May in the south, and well into June farther north (Palmer 1962, McIntyre 1975). Eggs are typically laid at two-day intervals (McIntyre 1975). Replacement clutches following failures of first nests are common (McIntyre 1975, 1988). Renests have been reported to occur within five days of a nest loss (Olson and Marshall 1952), but intervals of 10-14 days appear to be most common (Olson and Marshall 1952, McIntyre 1975, Sutcliffe 1980). Up to three laying cycles have been recorded in a season (Olson and Marshall 1952, McIntyre 1975). Nests lost early in the season are more likely to be replaced than those lost later (McIntyre 1988). Replacement clutches have been initiated as late as early July in Vermont (Kaveney and Rimmer 1989). If waters rise during incubation, loons continue adding to the nest's height to prevent flooding (McIntyre 1988). Replacement nests tend to have smaller outside dimensions (McIntyre 1975). Nest bowls are often reused in subsequent years, and occasionally within years for replacement clutches (Strong et al. 1987).
CLUTCH SIZE AND INCUBATION: Most clutches contain two eggs, and most one-egg clutches result from loss of the first egg (McIntyre 1975, Titus and VanDruff 1981). Three-egg clutches are very rare (Bent 1919, McIntyre 1988), and only two four-egg clutches have been reported (Nelson 1983, Zicus et al. 1983). Second eggs are smaller than first eggs, and eggs in replacement clutches are smaller than those in original clutches (McIntyre 1988). Both pair members incubate, beginning with the laying of the first egg, for an average period of 28-29 days, ranging from 26-31 days (Bent 1919, Olson and Marshall 1952, Palmer 1962, McIntyre 1975). An adult is present at the nest 99 percent of the time, and the eggs hatch within a day of one another (McIntyre 1975).
CHICK REARING: Chicks leave the nest within 24 hours of hatching and are soon moved to nursery areas (McIntyre 1988). In Saskatchewan, nurseries were located an average of 500 m from nest sites and occupied about 15 percent of territory size (McIntyre 1983). Both adults tend the young by feeding, carrying and defending them for several weeks. Chicks are carried on their parents' backs until they reach three weeks of age (McIntyre 1975). Although chicks are capable of short dives at the time of nest departure and may capture some fish by the second or third week (McIntyre 1975), they are fed largely by their parents until eight weeks of age (McIntyre 1988). Adults aggressively defend chicks underwater and on the surface (McIntyre 1988). Most juveniles are capable of flight at 11-12 weeks (Barr 1973, McIntyre 1975), and some leave their small, natal lakes or parental territories shortly afterwards (McIntyre 1975).
NESTING SUCCESS: Breeding success varies considerably among populations. Most failures occur during incubation, from factors such as predation, flooding or stranding due to water level fluctuations, and human intrusion (Olson and Marshall 1952, McIntyre 1975, Wood 1979, Titus and VanDruff 1981, Rimmer and Kaveney 1988). In Ontario, lack of attempted breeding was associated with small, brown, low-alkalinity lakes; successful breeding associated with large, clear, high-alkalinity lakes; unsuccessful breeding resulted primarily from brood mortalities on acidic lakes, most likely due to shotage of suitable food for young (Alvo et al. 1988).
Chick survival is relatively high, especially after chicks reach two to three weeks of age (McIntyre 1988). However, Alvo et al. (1988) recently found higher mortality of older chicks on highly acidified lakes in Ontario, due to presumed starvation from an inadequate food base. Fledging success (percent of hatched chicks fledged) from a sample of 1,500 pairs across the breeding range averaged 80 percent (range = 67-94 percent ) (McIntyre 1988). Productivity (number of fledglings per pair) of this sample averaged 0.60 and varied widely between 0.22 for nine pairs in Minnesota (McIntyre 1975) and 0.97 for 132 pairs in New York (Parker and Miller 1988).
SITE FIDELITY: Appear to be faithful to breeding territories. Banded adults have been recaptured on the same breeding territory in subsequent years (McIntyre 1974, Yonge 1981, Eberhardt 1984). Yearly reuse of nest sites and nursery areas has been documented (Strong et al. 1987, Jung 1991), but it is not known whether the same individuals were involved. Sonograms of yodel calls suggest that individual males return to the same territory each year (McIntyre 1988, Miller 1989). Little is known about mate fidelity of breeding pairs.