|Divers / Loons
Fossil range: ?Late Eocene - Recent (see text)
|The Pacific Loon (Gavia pacifica) is the sister species of the Black-throated Diver (G. arctica)|
Wetmore & Miller, 1926
The loons (North America) or divers (UK/Ireland) are a group of aquatic birds found in many parts of North America and northern Eurasia (Europe, Asia and debatably Africa). All living species of loons are members of one genus (Gavia), family (Gaviidae) and order (Gaviiformes) of their own.
The loons are the size of a large duck or small goose, which they somewhat resemble in shape when swimming. Like in these but unlike in coots (which are Rallidae) and grebes (Podicipedidae), their toes are connected by webbing. They may be confused even more readily with cormorants (Phalacrocoracidae), which are not too distant relatives of divers and like them are heaviset birds whose bellies – unlike those of ducks and geese – are submerged when swimming. Flying loons resemble a plump goose with a seagull's wings, which seem quite small in proportion to the bulky body. They hold their head slightly pointing upwards during swimming, less so than cormorants do, and in flight they let the head decidedly droop down compared to similar aquatic birds.
Males and females do not differ in plumage. Males are a bit larger on average, but usually this is only conspicuous when directly comparing the two parents. Their plumage is largely patterned black-and-white in summer, with grey on the head and neck in some species, and a white belly in all of them. This resembles many sea-ducks (Merginae) a lot – notably the smaller goldeneyes (Bucephala) – but is distinct from most cormorants which rarely have white feathers, and if so usually as large rounded patches rather than delicate patterns. All species of divers have a spear-shaped bill.
In winter plumage, they are dark gray above, with some indistinct lighter mottling on the wings, and a white chin, throat and underside. The species can then be distinguished by certain features, such as size and colour of head, neck, back and bill, but often reliable identification of wintering divers is tough even for experts – particularly as the smaller immature birds look similar to winter-plumage adults, making size also not fully reliable.
Gaviiformes are among the few groups of birds where the young moult into a second coat of down feathers after shedding the first one, rather than growing juvenile feathers with downy tips that wear off as is typical in birds. This trait is also found in tubenoses (Procellariiformes) and penguins (Sphenisciformes), presumably both quite close relatives of the loons.
Loons are excellent swimmers, using their feet to propel themselves above and under water and their wings for assistance. Because their feet are far back on the body, loons are poorly adapted to moving on land. They usually avoid going onto land, except when nesting.
All loons are decent fliers, though the larger species have some difficulty taking off and thus must swim into the wind to pick up enough velocity to become airborne. Only the Red-throated Diver (G. stellata) can take off from land. Once airborne, their considerable stamina allows them to migrate long distances southwards in winter, where they reside in coastal waters. Loons can live as long as 30 years.
Loons find their prey by sight. They eat mainly fish, supplemented with amphibians, crustaceans and similar mid-sized aquatic fauna. Specifically, they have been noted to feed on crayfish, frogs, snails, salamanders and leeches. They prefer clear lakes because they can see their prey more easily through the water. The loon uses its pointy bill to stab or grasp prey. They eat vertebrate prey headfirst to facilitate swallowing, and swallow all their prey whole.
To help digestion, loons swallow small pebbles from the bottoms of lakes. Similar to grit eaten by chickens, these gastroliths may assist the loon's gizzard in crushing the hard parts of the loon's food such as the exoskeletons of crustaceans and the bones of frogs and salamanders. The gastroliths may also be involved in stomach cleaning as an aid to regurgitation of indigestible food parts.
Loons may inadvertently ingest small lead pellets, released by anglers and hunters, which will slowly lead to the loon's death by lead poisoning. Jurisdictions that have banned the use of lead shot and sinkers include Maine, New Hampshire, Vermont, some areas of Massachusetts, Yellowstone National Park, Great Britain, Canada, and Denmark.
Gaviidae nest during the summer on freshwater lakes and/or large ponds. Smaller bodies of water (up to 0.5 km²) will usually only have one pair. Larger lakes may have more than one pair, with each pair occupying a bay or section of the lake.
Loons build their nests close to the water, preferring sites that are completely surrounded by water such as islands or emergent vegetation. Loons use a variety of materials to build their nests including aquatic vegetation, pine needles, leaves, grass, moss and mud. Both male and female build the nest and incubate jointly for 28 days. If the eggs are lost, the pair may re-nest, usually in a different location.
Despite the roughly equal participation of the sexes in nest building and incubation, analysis has shown clearly that males alone select the location of the nest. This pattern has the important consequence that male loons, but not females, establish significant site-familiarity with their territories that allows them to produce more chicks there over time. Sex-biased site-familiarity might explain, in part, why resident males fight so hard to defend their territories.
Most clutches consist of two eggs, which are laid in May or June, depending upon latitude. Loon chicks are precocial, able to swim and dive right away, but will often ride on their parents' back during their first 2 weeks to rest, conserve heat and avoid predators.
Chicks are fed mainly by their parents for about six weeks but gradually begin to feed themselves over time. By 11 or 12 weeks, chicks gather almost all of their own food and have learned to fly.
Biologists, especially from Chapman University, have extensively studied the mating behavior of the Common Loon (G. immer). Contrary to popular belief, pairs seldom mate for life. Indeed, a typical adult loon is likely to have several mates during its lifetime because of territorial takeover. Each breeding pair must frequently defend its territory against "floaters" (territory-less adults) trying to evict at least one owner and seize the breeding site. Territories that have produced chicks in the past year are especially prone to takeovers, because nonbreeding loons use chicks as cues to indicate high-quality territories. One-third of all territorial evictions among males result in the death of the owner; in contrast, female loons usually survive. Birds that are displaced from a territory but survive usually try to remate and (re)claim a breeding territory later in life.
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The European name "diver" comes from the bird's habit of catching fish by swimming calmly along the surface and then abruptly plunging into the water. The North American name "loon" probably originates from the identical Scots term for the Black-throated Diver. Altogether "loon" seem to be a reference to the bird's movements out of the water and/or its drawn-out calls. "Loon" seems to be an adoption of the Old Norse term lōmr, which may mean "lame ones". Related are words that express a ponderous, somewhat clumsy or lumbering quality – e.g. "lament" or the aforementioned "lame". Lomvie, another related term, means "Uria guillemot" – unrelated black-and-white seabirds that are also very clumsy on land – in most languages actually descended from Old Norse. The Scots word "loon" has also been influenced by the Middle English loun and as such may mean "crazy person". This and popular etymology influence of "lunatic" is also a likely reference to the birds' haunting, high-pitched yodelling cry, which is often heard in moonlit nights.
The scientific name Gavia was the Latin term for the Smew (Mergellus albellus). This small sea-duck is quite unrelated to loons and just happens to be another black-and-white seabird which swims and dives for fish. It is not likely that the Ancient Romans had much knowledge of loons, as these are limited to more northern latitudes and since the end of the last glacial period seem to have occurred only as rare winter migrants in the Mediterranean region.
The term gavia was transferred from the ducks to the loons only in the 18th century; earlier naturalists referred to the loons as mergus (the Latin term for diving seabirds of all sorts) or colymbus, which became the genus name used in the first modern scientific description of a Gavia species (by Carl Linnaeus) in 1758. Unfortunately, confusion about whether Linnaeus' "wastebin genus" Colymbus referred to loons or grebes abounded, with North American ornithologists using the genus name for grebes and European ones – following Nicholas Aylward Vigors and Richard Bowdler Sharpe – for divers. The International Commission on Zoological Nomenclature tried to settle this issue in 1956 by declaring Colymbus a suppressed name unfit for further use and establishing Gavia, created by Johann Reinhold Forster in 1788, as the valid genus name for the loons. However, the situation was not completely resolved even then, and the following year the ICZN had to act again to prevent Louis Jean Pierre Vieillot's 1818 almost-forgotten family name Urinatoridae from overruling the much younger Gaviidae. Nowadays, even though some eminent ornithologists such as Pierce Brodkorb tried to keep the debate alive at first, the ICZN's solution has been found very satisfactory.
The loons were formerly often considered to be the most ancient of the northern hemisphere bird families; this idea grew basically out of the perceived similarity of shape and (probably) habits between loons and the entirely unrelated extinct Cretaceous order Hesperornithiformes. In particular Enaliornis, which was apparently an ancestral and plesiomorphic member of that order, was sometimes used to support claims of Albian (Early Cretaceous) Gaviiformes.
More recently, it has become clear that the Anseriformes (waterfowl) and the Galliformes are the most ancient groups of modern birds, and these being distinct by the end of the Albian 100 million years ago (Ma), while just possible, is not at all well-supported. Loons belong to a more modern radiation. They were once believed to be related to grebes, which are also foot-propelled diving birds, and both species were once classified together under the order Colymbiformes. However, as recently as the 1930s, it was determined that the two groups are not that closely related at all and are merely the product of convergent evolution and adapted in a similar way to a similar ecological niche. The similarity is so strong that even the most modern cladistic analyses of general anatomical features are easily misled into grouping loons and grebes.
The Sibley-Ahlquist taxonomy still allied the loons with the grebes in its massively paraphyletic "Ciconiiformes", and it is almost certain that the relationships of loons lie with some of the orders placed therein. Namely, other recent authors have considered loons to share a rather close relationship with seabirds such as penguins (Sphenisciformes), tubenoses (Procellariiformes), waders (Charadriiformes) – and perhaps the newly-discovered clade Mirandornithes which unites grebes (Podicipediformes) and their closest living relatives, the flamingos (Phoenicopteriformes). It is perhaps notable that some early penguins had skulls and beaks that were in many aspects similar to those of the known living and fossil Gaviiformes.
In prehistoric times, the loons had a more southerly distribution than today, and their fossils have been found in places such as California, Florida and Italy. The conflicting molecular data regarding their relationships is not much resolved by the fossil record; though they seem to have originated at the end of the Late Cretaceous like their presumed relatives, modern loons are only known with certainty since the Eocene. By that time almost all modern bird orders are at least strongly suspected to have existed – if not known from unequivocally identified specimens – anyway.
Colymboides, the oldest unequivocal gaviiform genus known as of 2009, is widely known from early Priabonian – about 37 million years ago (Ma) in the Late Eocene – to Early Miocene (late Burdigalian, less than 20 Ma) limnic and marine rocks of western Eurasia north of the Alpide belt, between the Atlantic and the former Turgai Sea. It is usually placed in the Gaviidae already, but usually in a subfamily Colymboidinae, with the modern-type loons making up the Gaviinae. But the Colymboides material is generally quite distinct from modern loons, and may actually belong in a now-extinct family of primitive gaviiforms. Furthermore, the supposed genus could well be paraphyletic, so that for example Dyspetornis – which is now contained therein – might have to be separated again. A leg of an undescribed small diver was found in the Late Oligocene deposits at Enspel (Germany); it too may or may not belong to Colymboides. Of the crown genus Gavia, nearly ten prehistoric species have been named to date, and about as many undescribed ones await further study. The genus is known from the Early Miocene onwards, and the oldest members them are rather small (some are smaller than the Red-throated Diver). Throughout the late Neogene, the genus by and large follows Cope's Rule.
List of fossil Gavia
"Gavia" portisi from the Late Pliocene of Orciano Pisano (Italy) is known from a cervical vertebra that may or may not have been from a loon. If so, it was from a bird slightly smaller than Common Loon. Older authors were quite sure the bone was indeed from a Gavia and even considered G. concinna a possibly junior synonym of it. This is now regarded as rather unlikely due to the quite distinct range and age. Interestingly, the Early Pliocene Gavia skull from Empoli (Italy) was referred to G. concinna, and thus could conceivably have been of "G." portisi if that was indeed a diver. The holotype vertebra may now be lost, which would make "G." portisi a nomen dubium.
In addition, there are some older fossils that are sometimes assigned to the Gaviiformes. From the Late Cretaceous, the genera Lonchodytes (Lance Formation, Wyoming) and Neogaeornis (Quinriquina Formation, Chile) have been described; both are usually allied with orders which are considered related to loons. In particular the latter is still sometimes explicitly proposed as a primitive loon as they both were initially, but other authors consider Neogaeornis a hesperornithiform; note however that neither Gaviiformes nor Hesperornithiformes are known from the Southern Hemisphere or anywhere near it. Lonchodytes was more certainly quite close to loons, but probably closer still to some of the loons' relatives. Of similarly doubtful validity and surrounded by considerable dispute is the supposed Late Cretaceous loon Polarornis (Seymour Island, Antarctica). Eupterornis from the Paleocene of Châlons-sur-Vesle (France) has some features reminiscent of loons, but others seem more similar to Charadriiformes such as gulls (Laridae). A piece of a carpometacarpus supposedly from Oligocene rocks near Lusk, Wyoming was described as Gaviella pusilla, but this handbone also shows some similarities to the plotopterids which were flightless wing-propelled divers and if these are apomorphic would make an unconvincing member of the Gaviidae (though it still could be a small-winged gaviiform in an – as of yet undescribed – family "Gaviellidae"): while the carpometacarpus in Gavia is somewhat convergent to that of wing-propelled divers, enabling the wings to be used as rudders for quick underwater turns, Colymboides still had an unspecialized plesiomorphic hand. Parascaniornis, sometimes allied to the loons by early authors, was eventually determined to be a junior synonym of the hesperornithiform Baptornis. A supposed mid-Eocene diver fossil form Geiseltal (Germany) was erroneously assigned to Gavia.