Loon

Divers / Loons Fossil range: ?Late Eocene - Recent (see text) 37–0 Ma PreЄ Є O S D C P T J K Pg N
The Pacific Loon (Gavia pacifica) is the sister species of the Black-throated Diver (G. arctica)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Subclass:	Neornithes
Infraclass:	Neognathae
Order:	Gaviiformes Wetmore & Miller, 1926
Family:	Gaviidae Coues, 1903
Genus:	Gavia Forster, 1788
Diversity
5 species
Synonyms
Order-level: Colymbiformes Sharpe, 1891 (but see text) Family-level: Colymbidae Vigors, 1825 (but see text) Colymbinae Bonaparte, 1831 (but see text) Urinatores Vieillot, 1818 Urinatoridae Vieillot, 1818 Urinatorides Vieillot, 1818 Genus-level: Colymbus Linnaeus, 1758 (but see text) Urinator Lacépède, 1799

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 the genus (Gavia), family (Gaviidae) and order (Gaviiformes).

Description

Typical flight profile of a Gavia species

Great Northern Diver or Common Loon (G. immer) rearing up. Note the plump body and pointed but rather short wings.

The loon is the size of a large duck or small goose, which it resembles in shape when swimming. Like ducks and geese but unlike coots (which are Rallidae) and grebes (Podicipedidae), the loon's toes are connected by webbing. The bird may be confused with cormorants (Phalacrocoracidae), not too distant relatives of divers and like them are heavy set birds whose bellies – unlike those of ducks and geese – are submerged when swimming. Flying loons resemble a plump goose with a seagull's wings, relatively small in proportion to the bulky body. The bird holds its head pointing slightly upwards during swimming, but less so than cormorants do. In flight the head droops more than in similar aquatic birds.

Male and female loons have identical plumage. 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. 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.

Males are larger on average, but only apparent when the male and female are together.

In winter plumage is 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 way for identification.^[1]

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 many birds. This trait is also found in tubenoses (Procellariiformes) and penguins (Sphenisciformes), presumably both quite close relatives of the loons.^[2]

Ecology

Loons on Wolfe Lake (Ontario, Canada)

Loons are excellent swimmers, using their feet to propel themselves above and under water while their wings provide assistance. Because their feet are far back on the body, loons are poorly adapted to moving on land, and 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.

Food and feeding

Common Loon feeding its young

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 more easily see their prey 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, that will contribute to lead poisoning and the loon's eventual death. Jurisdictions that have banned the use of lead shot and sinkers include but are not limited to Maine, New Hampshire, Vermont, some areas of Massachusetts, Yellowstone National Park, Great Britain, Canada, and Denmark.^[3]

Reproduction

Juvenile Red-throated Diver

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.^[4]

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.^[5]

Etymology and taxonomy

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^[6] 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.^[7]

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.^[8]

Systematics and evolution

All living species are placed in the genus Gavia. Phylogenetically, the 5 living species can be arranged as follows:^[9]

White-billed Diver or Yellow-billed Loon (Gavia adamsii) in winter plumage

Basal lineage

• Red-throated Loon or Red-throated Diver, Gavia stellata

Black-throated lineage

• Black-throated Loon, Arctic Loon or Black-throated Diver, Gavia arctica
• Pacific Loon or Pacific Diver, Gavia pacifica (formerly in G. arctica)

Black-headed lineage

• Great Northern Loon, Common Loon or Great Northern Diver, Gavia immer
• Yellow-billed Loon or White-billed Diver, Gavia adamsii

Relationships

The Paleocene penguin ancestor Waimanu resembled loons in some aspects of its head and bill, but it was already flightless and used its feet for steering rather than propulsion

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.^[10]

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.^[11]

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.^[12]

Fossil record

Red-throated Diver or Red-throated Loon (G. stellata), the smallest living Gavia species. Some Miocene members of this genus were smaller still.

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.^[13]

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^[14] 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.^[15]

List of fossil Gavia

• Gavia sp. (Early-Middle Miocene of E USA)^[16]
• Gavia egeriana Švec, 1982 (Early Miocene of Czechoslovakia ?and Cheswold, Delaware, USA –? Yorktown Early Pliocene of Lee Creek Mine, South Carolina, USA)^[17]
• Gavia schultzi Mlíkovský, 1998 (Middle Miocene of Sankt Margarethen, Austria)^[18]
• Gavia sp. (Calvert Middle Miocene ?or Pleistocene of Maryland, USA) – same as Gavia cf. immer below?^[19]
• Gavia spp. (Middle Miocene of Steinheim, Germany) – three species^[18]
• Gavia brodkorbi Howard, 1978 (Late Miocene of Orange County, USA)^[20]
• Gavia moldavica Kessler, 1984 (Late Miocene of Chişinău, Moldova)^[18]
• Gavia paradoxa Umanska, 1981 (Late Miocene of Čebotarevka, Ukraine)^[18]
• Gavia concinna Wetmore, 1940 (Late Miocene/Early Pliocene of W and E USA)^[21]
• Gavia fortis Olson & Rasmussen, 2001 (Yorktown Early Pliocene of Lee Creek Mine, South Carolina, USA)
• Gavia sp. (Early Pliocene of Empoli, Italy)^[22]
• Gavia sp. (Early Pliocene of Kerč Peninsula, Ukraine)^[18]
• Gavia palaeodytes Wetmore, 1943 (Bone Valley Early/Middle Pliocene of Pierce, Florida, USA)^[23]
• Gavia howardae Brodkorb, 1953 (Yorktown Early Pliocene of Lee Creek Mine, South Carolina, USA – San Diego Middle Pliocene of San Diego, California, USA)^[24]
• Gavia cf. concinna (San Diego Middle/Late Pliocene of San Diego, California, USA) – two species?^[25]
• Gavia sp. (Early Pleistocene of Kairy, Ukraine)^[18]
• Gavia cf. immer (Pleistocene of California and Florida, USA) – possibly a G. immer paleosubspecies^[26]

"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.^[27]

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^[28] 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"^[29]): 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.^[30]

Loons in popular culture

• The Common Loon is the provincial bird of Ontario and is depicted on the Canadian one-dollar coin, which has come to be known affectionately as the "loonie".^[31]
• It is also the official state bird of Minnesota.^[32]
• Mercer, Wisconsin promotes itself as the "Loon Capital of the World".^[33]
• Loons are mentioned several times in the Mark Rydell/Ernest Thompson film On Golden Pond and the Stephen King novels Pet Semetary and Gerald's Game.
• The Common or Great Northern Loon features in the Arthur Ransome children's book Great Northern?.
• The loon is mentioned in the Nick Cave and The Bad Seeds song "Lime Tree Arbour".

Footnotes

References

• Appleby, R.H.; Steve C. Madge & Mullarney, Killian (1986): Identification of divers in immature and winter plumages. British Birds 79(8): 365-391.
• Arnott, W.G. (1964): Notes on Gavia and Mergvs in Latin Authors. Classical Quarterly (New Series) 14(2): 249-262. First page image
• Brodkorb, Pierce (1953): A Review of the Pliocene Loons. Condor 55(4): 211-214. DjVu fulltext PDF fulltext
• Brodkorb, Pierce (1963): Catalogue of fossil birds. Part 1 (Archaeopterygiformes through Ardeiformes). Bulletin of the Florida State Museum, Biological Sciences 7(4): 179-293. PDF fulltext
• Heinrichs, Ann (2003): Minnesota. Compass Point Books. ISBN 0756503159
• International Commission on Zoological Nomenclature (ICZN) (1957–58): The family-group names "Gaviidae" Coues, 1903 and "Urinatoridae" (correction of "Urinatores)" [sic] Vieillot, 1818 (Class Aves) – "Opinion" 401 and "Direction" 75. Bulletin of Zoological Nomenclature 15A: 147-148. Fulltext at the Internet Archive
• Linnaeus, Carl (1758): 68.1. Colymbus arcticus. In: Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (10th ed., vol. 1): 190 [Latin book]. Lars Salvius, Stockholm ("Holmius"). Digitized version
• Mayr, Gerald (2004): A partial skeleton of a new fossil loon (Aves, Gaviiformes) from the early Oligocene of Germany with preserved stomach content. J. Ornithol. 145: 281-286 doi:10.1007/s10336-004-0050-9 PDF fulltext
• Mayr, Gerald (2009): Paleogene Fossil Birds. Springer-Verlag, Heidelberg & New York. ISBN 3540896279
• Minnesota Pollution Control Agency (2007): Let's Get the Lead Out! Non-lead alternatives for fishing tackle. Version of June, 2007. Retrieved 2007-JUL-23.
• Mlíkovský, Jirí (2002): Cenozoic Birds of the World, Part 1: Europe. Ninox Press, Prague. ISBN 80-901105-3-8 PDF fulltext
• Montana Fish, Wildlife & Parks (Montana FW&P) (2007): Animal Field Guide: Common Loon. Retrieved 2007-MAY-12.
• Moran, Mark; Sceurman, Mark; Godfrey, Linda S. & Hendricks, Richard D. (2005): Weird Wisconsin: Your Travel Guide to Wisconsin's Local Legends and Best Kept Secrets. Sterling Publishing. ISBN 0760759448
• Olson, Storrs L. (1985): Section X.I. Gaviiformes. In: Farner, D.S.; King, J.R. & Parkes, Kenneth C. (eds.): Avian Biology 8: 212-214. PDF fulltext
• Piper, W.H.; Evers, D.C.; Meyer, M.W.; Tischler, K.B. & Klich, M. (2000a): Do common loons mate for life?: scientific investigation of a widespread myth. In: McIntyre, J. & Evers, D.C. (eds.): Loons: old history and new findings – proceedings of a symposium from the 1997 meeting of the American Ornithologists' Union: 43-49. North American Loon Fund, Meredith, New Hampshire.
• Piper, W.H.; Tischler, K B. & Klich, M. (2000b): Territory acquisition in loons: the importance of take-over. Animal Behaviour 59(2): 385-394. doi:10.1006/anbe.1999.1295 (HTML abstract)
• Piper, W.H.; Walcott, C.; Mager, J.N.; Perala, M.; Tischler, K B.; Harrington, Erin; Turcotte, A. J.; Schwabenlander M. & Banfield, N. (2006): Prospecting in a Solitary Breeder: Chick Production Elicits Territorial Intrusions in Common Loons. Behavioral Ecology 17(6): 881-888. doi:10.1093/beheco/arl021 PDF fulltext
• Piper, W.H.; Walcott, C.; Mager, J.N. & Spilker, F. (2008a): Nestsite selection by male loons leads to sex-biased site familiarity. Journal of Animal Ecology 77(2): 205-210. doi:10.1111/j.1365-2656.2007.01334.x (HTML abstract)
• Piper, W.H.; Walcott, C.; Mager, J.N. & Spilker, F. (2008b): Fatal Battles in Common Loons: A Preliminary Analysis. Animal Behaviour 75(3): 1109-1115. doi:10.1016/j.anbehav.2007.10.025 (HTML abstract)
• Rasmussen, Pamela C. (1998): Early Miocene Avifauna from the Pollack Farm Site, Delaware. Delaware Geological Survey Special Publication 21: 149-151. PDF fulltext
• Slack, K.E.; Jones, C.M.; Ando, T.; Harrison G.L.; Fordyce R.E.; Arnason, U. & Penny, D. (2006): Early Penguin Fossils, plus Mitochondrial Genomes, Calibrate Avian Evolution. Mol. Biol. Evol. 23(6): 1144-1155. doi:10.1093/molbev/msj124 PDF fulltext Supplementary Material
• Stewart, Barry D. (2004): Across The Land: A Canadian Journey Of Discovery. Trafford Publishing. ISBN 1412022762
• Stolpe, M. (1935): Colymbus, Hesperornis, Podiceps:, ein Vergleich ihrer hinteren Extremität. J. Ornithol. 80(1): 115-128. [Article in German] doi:10.1007/BF01908745
• Storer, Robert W. (1956): The Fossil Loon, Colymboides minutus. Condor 58(6): 413-426. DjVu fulltext PDF fulltext
• Wetmore, Alexander (1941): An Unknown Loon from the Miocene Fossil Beds of Maryland. Auk 58(4): 567. DjVu fulltext PDF fulltext
• United States Fish and Wildlife Service (USFWS) (2005): Common Loons at Seney NWR, June 2005. PDF fulltext
• Wings, Oliver (2007): A review of gastrolith function with implications for fossil vertebrates and a revised classification. Acta Palaeontologica Polonica 52: 1-16 PDF fulltext

External links

• Video of nesting loons. Video captures the laying and hatching of two eggs on Lake George in Minnesota.Schara, Ron: The Life of a Loon. KARE11. Version of 2007-MAY-15. Retrieved 2007-NOV-10.
• Tree of Life Gaviidae
• Loon sounds
• Loon videos on the Internet Bird Collection
• Loon Preservation Committee
• Diving Birds of North America, by Paul Johnsgard

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