|Black-necked grebe (Podiceps nigricollis nigricollis), in non-breeding plumage|
Grebes are a widely distributed order of freshwater diving birds, some of which visit the sea when migrating and in winter. This order contains only a single family, the Podicipedidae, containing 22 species in six extant genera.
Grebes are small to medium-large in size, have lobed toes, and are excellent swimmers and divers. Although they can run for a short distance, they are prone to falling over, since they have their feet placed far back on the body.
Grebes have narrow wings, and some species are reluctant to fly; indeed, two South American species are completely flightless. They respond to danger by diving rather than flying, and are in any case much less wary than ducks. Extant species range in size from the least grebe, at 120 grams (4.3 oz) and 23.5 cm (9.3 inches), to the great grebe, at 1.7 kg (3.8 lbs) and 71 cm (28 inches).
The North American and Eurasian species are all, of necessity, migratory over much or all of their ranges, and those species that winter at sea are also seen regularly in flight. Even the small freshwater pied-billed grebe of North America has occurred as a transatlantic vagrant to Europe on more than 30 occasions.
Bills vary from short and thick to long and pointed, depending on the diet, which ranges from fish to freshwater insects and crustaceans. The feet are always large, with broad lobes on the toes and small webs connecting the front three toes. The hind toe also has a small lobe. Recent experimental work has shown that these lobes work like the hydrofoil blades of a propeller. Curiously, the same mechanism apparently evolved independently in the extinct Cretaceous-age Hesperornithiformes, which are totally unrelated birds.
Grebes have unusual plumage. It is dense and waterproof, and on the underside the feathers are at right-angles to the skin, sticking straight out to begin with and curling at the tip. By pressing their feathers against the body, grebes can adjust their buoyancy. Often, they swim low in the water with just the head and neck exposed. They swim by simultaneously spreading out the feet and bring them inward with the webbing expanded to produce the forward thrust in much the same way as frogs.
In the non-breeding season, grebes are plain-coloured in dark browns and whites. However, most have ornate and distinctive breeding plumages, often developing chestnut markings on the head area, and perform elaborate display rituals. The young, particularly those of the genus Podiceps, are often striped and retain some of their juvenile plumage even after reaching full size. In the breeding season, they mate at freshwater lakes and ponds, but some species spend their non-breeding season along seacoasts.
When preening, grebes eat their own feathers, and feed them to their young. The function of this behaviour is uncertain but it is believed to assist with pellet formation, and to reduce their vulnerability to gastric parasites.
Taxonomy, systematics and evolution
The grebes are a radically distinct group of birds as regards their anatomy. Accordingly, they were at first believed to be related to the loons, which are also foot-propelled diving birds, and both families were once classified together under the order Colymbiformes. However, as recently as the 1930s, this was determined to be an example of convergent evolution by the strong selective forces encountered by unrelated birds sharing the same lifestyle at different times and in different habitat. Grebes and loons are now separately classified orders of Podicipediformes and Gaviiformes, respectively.
The cladistics vs. phenetics debate of the mid-20th century revived scientific interest in generalizing comparisons. As a consequence, the discredited grebe-loon link was discussed again. This even went as far as proposing monophyly for grebes, loons, and the toothed Hesperornithiformes. In retrospect, the scientific value of the debate lies more in providing examples that a cladistic methodology is not incompatible with an overall phenetical scientific doctrine, and that thus, simply because some study "uses cladistics", it does not guarantee superior results.
Molecular studies such as DNA-DNA hybridization (Sibley & Ahlquist, 1990)[full citation needed] and sequence analyses fail to resolve the relationships of grebes properly due to insufficient resolution in the former and long-branch attraction in the latter. Still – actually because of this – they do confirm that these birds form a fairly ancient evolutionary lineage (or possibly one that was subject to selective pressures down to the molecular level even), and they support the non-relatedness of loons and grebes.
Relationship with flamingos
Recent molecular studies have suggested a relation with flamingos while morphological evidence also strongly supports a relationship between flamingos and grebes. They hold at least eleven morphological traits in common, which are not found in other birds. Many of these characteristics have been previously identified in flamingos, but not in grebes. The fossil Palaelodids can be considered evolutionarily, and ecologically, intermediate between flamingos and grebes.
For the grebe-flamingo clade, the taxon Mirandornithes ("miraculous birds" due to their extreme divergence and apomorphies) has been proposed. Alternatively, they could be placed in one order, with Phoenocopteriformes taking priority.
The fossil record of grebes is incomplete; there are no transitional forms between more conventional birds and the highly derived grebes known from fossils, or at least none that can be placed in the relationships of the group with any certainty. The enigmatic waterbird genus Juncitarsus, however, may be close to a common ancestor of flamingos and grebes.
The Early Cretaceous (Berriasian, around 143 mya) genus Eurolimnornis from NW Romania was initially believed to be a grebe. If it is indeed related to this lineage, it must represent a most basal form, as it almost certainly predates any grebe-flamingo split. On the other hand, the single bone fragment assigned to this taxon is not very diagnostic and may not be of a bird at all.
Telmatornis from the Navesink Formation – also Late Cretaceous – is traditionally allied with the Charadriiformes and/or Gruiformes. However, a cladistic analysis of the forelimb skeleton found it highly similar to the great crested grebe and unlike the painted buttonquail (now known to be a basal charadriiform lineage), the black-necked stilt (a more advanced charadriiform), or the limpkin (a member of the Grui suborder of Gruiformes), namely in that its dorsal condyle of the humerus was not angled at 20°–30° away from long axis of the humerus. The analysis did not result in a phylogenetic pattern but rather grouped some birds with similar wing shapes together while others stood separate. It is thus unknown whether this apparent similarity to grebes represents an evolutionary relationship, or whether Telmatornis simply had a wing similar to that of grebes and moved it like they do.
True grebes suddenly appear in the fossil record in the Late Oligocene or Early Miocene, around 23–25 mya. While there are a few prehistoric genera that are now completely extinct; Thiornis (Late Miocene -? Early Pliocene of Libros, Spain) and Pliolymbus (Late Pliocene of WC USA – Early? Pleistocene of Chapala, Mexico) date from a time when most if not all extant genera were already present. Because grebes are evolutionarily isolated and they only started to appear in the Northern Hemisphere fossil record in the Early Miocene, they are likely to have originated in the Southern Hemisphere.
A few more recent grebe fossils could not be assigned to modern or prehistoric genera:
- Podicipedidae gen. et sp. indet. (San Diego Late Pliocene of California) – formerly included in Podiceps parvus
- Podicipedidae gen. et sp. indet. UMMP 49592, 52261, 51848, 52276, KUVP 4484 (Late Pliocene of WC USA)
- Podicipedidae gen. et sp. indet. (Glenns Ferry Late Pliocene/Early Pleistocene of Idaho, USA)
Living Podicipediformes based on the work by John Boyd.
- Genus †Miobaptus Švec 1982
- Genus †Miodytes Dimitreijevich, Gál & Kessler 2002
- †Miodytes serbicus Dimitreijevich, Gál & Kessler 2002
- Genus †Pliolymbus Murray 1967 [Piliolymbus (sic)]
- †Pliolymbus baryosteus Murray 1967
- Genus †Thiornis Navás 1922
- †Thiornis sociata Navás 1922 [Podiceps sociatus (Navás 1922) Olson 1995]
- Genus Podilymbus Lesson 1831
- †P. mujusculus Murray 1967
- †P. wetmorei Storer 1976
- †Atitlán grebe, Podilymbus gigas Griscom 1929 (extinct 1989)
- Pied-billed grebe, Podilymbus podiceps (Linnaeus 1758)
- †P. p. magnus
- P. p. antillarum Bangs 1913 (Antillean pied-billed grebe)
- P. p. podiceps (Linnaeus 1758) (northern pied-billed grebe)
- P. p. antarcticus (Lesson 1842) (southern pied-billed grebe)
- Genus Tachybaptus Reichenbach 1853 [Poliocephalus (Tachybaptus) Reichenbach 1853; Sylbeocyclus Macgillivray 1842 non Bonaparte 1831; Limnodytes Oberholser 1974]
- Little grebe, Tachybaptus ruficollis (Pallas 1764)
- T. r. ruficollis (Pallas 1764) (European little grebe)
- T. r. albescens (Blanford 1877) (Indian little grebe)
- T. r. iraquensis (Ticehurst 1923) (Iraq little grebe)
- T. r. capensis (Salvadori 1884) (African little grebe)
- T. r. poggei (Reichenow 1902)
- T. r. philippensis (Bonnaterre 1790) (Philippine little grebe)
- T. r. cotabato (Rand 1948) (Mindanao little grebe)
- Tricolored grebe, Tachybaptus tricolor (Gray 1861)
- T. t. vulcanorum (Rensch 1929)
- T. t. tricolor (Gray 1861)
- T. t. collaris (Mayr 1945)
- Australasian grebe Tachybaptus novaehollandiae (Stephens 1826)
- T. n. javanicus (Mayr 1943)
- T. n. fumosus (Mayr 1943)
- T. n. incola (Mayr 1943)
- T. n. novaehollandiae (Stephens 1826) (Australian little grebe)
- T. n. leucosternos (Mayr 1931)
- T. n. rennellianus (Mayr 1943)
- Madagascar grebe, Tachybaptus pelzelnii (Hartlaub 1861)
- †Alaotra grebe, Tachybaptus rufolavatus (Delacour 1932) (extinct 2010)
- Least grebe, Tachybaptus dominicus (Linnaeus 1766)
- T. d. brachypterus (Chapman 1899) (Mexican least grebe)
- T. d. bangsi (van Rossem & Hachisuka 1937)] (Bangs' grebe)
- T. d. dominicus (Linnaeus 1766) (West Indian grebe)
- T. d. brachyrhynchus (Chapman 1899) (short-billed grebe)
- T. d. eisenmanni Storer & Getty 1985
- Little grebe, Tachybaptus ruficollis (Pallas 1764)
- Genus Poliocephalus Selby 1840 (Hoary-headed Grebe)
- Genus Aechmophorus Coues 1862)
- Genus Podicephorus Bochenski 1994
- Great grebe, Podicephorus major (Boddaert 1783) Bochenski 1994
- P. m. major (Boddaert 1783)
- P. m. navasi Manghi 1984
- Great grebe, Podicephorus major (Boddaert 1783) Bochenski 1994
- Genus Podiceps Latham 1787 [Pliodytes Brodkorb 1953; Dyas; Lophaithyia Kaup 1829; Colymbus Linnaeus 1758 non Linnaeus 1766 non Paetel 1875 non Hadding 1913; Podiceps (Proctopus) Kaup.; Centropelma Sclater & Salvin 1869; Dytes Kaup.; Rollandia Bonaparte, 1856]
- †P. oligocaenus (Shufeldt 1915)
- †P. arndti Chandler 1990
- †P. caspicus (Habizl 1783) [Colymbus caspicus Habizl 1783]
- †P. csarnotatus Kessler 2009
- †P. discors Murray 1967
- †P. dixi Brodkorp 1963
- †P. miocenicus Kessler 1984
- †P. oligocaenus (Shufeldt)
- †P. parvus (Shufeldt 1913)) [Colymbus parvus Schufeldt 1913]
- †P. solidus Kuročkin 1985
- †P. subparvus (Miller & Bowman 1958) [Colymbus subparvus Miller & Bowman 1958]
- White-tufted grebe, Podiceps rolland Quoy & Gaimard 1824
- P. r. rolland Quoy & Gaimard 1824 (Falkland white-tufted Grebe)
- P. r. chilensis Lesson 1828 (Chilean white-tufted Grebe)
- P. r. morrisoni Simmons 1962 (Junín white-tufted grebe)
- Titicaca grebe, Podiceps microptera Gould 1868
- Red-necked grebe, Podiceps grisegena (Boddaert 1783)
- P. g. grisegena (Boddaert 1783) (gray-cheeked grebe)
- P. g. holbollii Reinhardt 1853 (Holbøll's grebe)
- Great crested grebe, Podiceps cristatus (Linnaeus 1758)
- P. c. cristatus (Linnaeus 1758) (Eurasian great crested grebe)
- P. c. infuscatus Salvadori 1884 (African great crested grebe)
- P. c. australis Gould 1844 (Australasian great crested grebe)
- Horned grebe or Slavonian grebe, Podiceps auritus (Linnaeus 1758)
- P. a. auritus (Linnaeus 1758) (Eurasian Horned Grebe)
- P. a. cornutus (Gmelin 1789)
- Black-necked grebe or eared grebe, Podiceps nigricollis Brehm 1831
- P. n. nigricollis Brehm 1831 (Eurasian Black-necked Grebe)
- P. n. gurneyi (Roberts 1919) (African black-necked grebe)
- †Colombian grebe, Podiceps andinus (Meyer de Schauensee 1959) (extinct 1977)
- Silvery grebe, Podiceps occipitalis Garnot 1826
- P. o. juninensis von Berlepsch & Stolzmann 1894 (northern silvery grebe)
- P. o. occipitalis Garnot 1826 (Southern silvery Grebe)
- Junin grebe, Podiceps taczanowskii von Berlepsch & Stolzmann 1894
- Hooded grebe, Podiceps gallardoi Rumboll 1974
- North-American Eared grebe, Podiceps californicus Heermann 1854
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