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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Ordo: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Euavialae
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Cladus: Neoaves
Cladus: Aequornithes
Ordo: Gaviiformes

‎Infraclassis: Aves
Cohors: Neognathae
Cladus: Neoaves
Cladus: Aequornithes
Ordo: Gaviiformes
Familiae: Gaviidae
Genera incertae sedis: †Petralca

Gaviiformes Wetmore & Miller, 1926: 340
Primary references

Wetmore, A. & Miller, W.D. 1926. The revised classification for the fourth edition of the A. O. U. check-list. The Auk 43(3): 337–346. DOI: 10.2307/4075427. Reference page.


Gaviiformes – Taxon details on Fossilworks.
Kagu, Sunbittern, tropicbirds, loons, penguins – IOC World Bird List v12.1

Vernacular names
Deutsch: Seetaucher
Esperanto: Gavioformaj
Nordfriisk: Siadükerfögler
magyar: Búváralakúak
日本語: アビ目
македонски: Нуркачовидни
Nederlands: Zeeduikerachtigen
polski: Nury
slovenščina: Slapniki
Türkçe: Dalgıç kuşları
українська: Гагароподібні
中文: 潜鸟目

Gaviiformes is an order of aquatic birds containing the loons or divers and their closest extinct relatives. Modern gaviiformes are found in many parts of North America and northern Eurasia (Europe, Asia and debatably Africa), though prehistoric species were more widespread.

Classification and evolution
The Paleocene Waimanu (an early member of the Sphenisciformes) resembled loons in some aspects of its head and bill, but it was already flightless and used its feet for steering rather than propulsion

There are five living species, and all are placed in the genus Gavia.[2] 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.[3][4]

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 groups 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.[3][5][6]

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.[5][7]
Fossil record
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.[6]

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[8] 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 of them are rather small (some are smaller than the red-throated loon). Throughout the late Neogene, the genus by and large follows Cope's Rule.[9]

Some older fossils 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. 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 a yet-undescribed family "Gaviellidae"[10]): 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.[11]
See also

List of Gaviiformes by population


Kuhl., H.; Frankl-Vilches, C.; Bakker, A.; Mayr, G.; Nikolaus, G.; Boerno, S. T.; Klages, S.; Timmermann, B.; Gahr, M. (2020). "An unbiased molecular approach using 3'UTRs resolves the avian family-level tree of life". Molecular Biology and Evolution: 143. doi:10.1093/molbev/msaa191.
Boertmann, D. (1990). "Phylogeny of the divers, family Gaviidae (Aves)". Steenstrupia. 16: 21–36.
Stolpe, M. (January 1935). "Colymbus, Hesperornis, Podiceps:, ein Vergleich ihrer hinteren Extremität". J. Ornithol. (in German). 83 (1): 115–128. doi:10.1007/BF01908745.
Brodkorb (1963: pp. 220–221)
Slack, K.E.; Jones, C.M.; Ando, T.; Harrison G.L.; Fordyce R.E.; Arnason, U.; Penny, D. (June 2006). "Early Penguin Fossils, plus Mitochondrial Genomes, Calibrate Avian Evolution". Mol. Biol. Evol. 23 (6): 1144–1155. doi:10.1093/molbev/msj124. PMID 16533822. Supplementary Material Archived 2009-12-16 at the Wayback Machine
Mayr (2009)
Olson (1985: pp. 212–213), Mayr (2004, 2009)
Some (notably Robert W. Storer) have disagreed, usually because they separated Gaviella in the basalmost subfamily of the Gaviidae and considered Colymboides the ancestor of Gavia. More recent authors generally disagree at least regarding the latter: Storer (1956), Olson (1985), Mayr (2009: pp. 75–76)
Brodkorb (1953), 1963: pp. 223–225, Olson (1985: pp. 212–213), Mlíkovský (2002: pp. 63–64), Mayr (2009: pp. 75–76)
Not to be used without quotation marks, as it is not a valid taxon.

Brodkorb (1963: pp. 220–223), Olson (1985), Mlíkovský (2002: pp. 64, 259–261), Mayr (2009: p. 20)


Brodkorb, Pierce (1953). "A Review of the Pliocene Loons" (PDF). Condor. 55 (4): 211–214. doi:10.2307/1364769. JSTOR 1364769.
Brodkorb, Pierce (1963). "Catalogue of fossil birds. Part 1 (Archaeopterygiformes through Ardeiformes)". Bulletin of the Florida State Museum, Biological Sciences. 7 (4): 179–293.
Mayr, Gerald (2004). "A partial skeleton of a new fossil loon (Aves, Gaviiformes) from the early Oligocene of Germany with preserved stomach content" (PDF). J. Ornithol. 145 (4): 281–286. doi:10.1007/s10336-004-0050-9. Archived from the original (PDF) on 2015-09-24. Retrieved 2011-01-14.
Mayr, Gerald (2009). Paleogene Fossil Birds. Heidelberg & New York: Springer-Verlag. ISBN 978-3-540-89627-2.
Mlíkovský, Jirí (2002). Cenozoic Birds of the World, Part 1: Europe (PDF). Ninox Press, Prague.
Olson, Storrs L. (1985). "Section X.I. Gaviiformes" (PDF). In Farner, D.S.; King, J.R.; Parkes, Kenneth C. (eds.). Avian Biology. Vol. 8. pp. 212–214.
Storer, Robert W. (1956). "The Fossil Loon, Colymboides minutus" (PDF). Condor. 58 (6): 413–426. doi:10.2307/1365096. JSTOR 1365096.

Further reading
Appleby, R.H.; Madge, Steve C.; 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. doi:10.1017/S0009838800023806. JSTOR 637729.
Heinrichs, Ann (2003): Minnesota. Compass Point Books. ISBN 0-7565-0315-9
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.
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 Archived 2017-06-13 at the Wayback Machine
Minnesota Pollution Control Agency (2007): Let's Get the Lead Out! Non-lead alternatives for fishing tackle. Version of June, 2007. Retrieved 2007-July-23.
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 0-7607-5944-8
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. PMID 10675261.
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.
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. PMID 17976165.
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.
Rasmussen, Pamela C. (1998). "Early Miocene Avifauna from the Pollack Farm Site, Delaware". Delaware Geological Survey Special Publication. 21: 149–151.
Stewart, Barry D. (2004): Across The Land: A Canadian Journey Of Discovery. Trafford Publishing. ISBN 1-4120-2276-2
Wetmore, Alexander (1941). "An Unknown Loon from the Miocene Fossil Beds of Maryland" (PDF). Auk. 58 (4): 567. doi:10.2307/4078641. JSTOR 4078641.
United States Fish and Wildlife Service (USFWS) (2005): Common Loons at Seney NWR, June 2005.
Wings, Oliver (2007). "A review of gastrolith function with implications for fossil vertebrates and a revised classification" (PDF). Acta Palaeontologica Polonica. 52: 1–16. Archived from the original (PDF) on 2008-03-07.

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