Grebe
100005
225147544
2008-07-12T03:34:34Z
Nicolás10
5633280
{{Taxobox
| name = Grebes
| image =Aechmophorus_clarkii_on_nest.jpg
| image_width = 250px
| image_caption = [[Clark's Grebe]], ''Aechmophorus clarkii''
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Bird|Aves]]
| infraclassis = [[Neognathae]]
| ordo = '''Podicipediformes'''
| ordo_authority = [[Max Fürbringer|Fürbringer]], 1888
| familia = '''Podicipedidae'''
| familia_authority = [[Charles Lucien Jules Laurent Bonaparte|Bonaparte]], 1831
| subdivision_ranks = Genera
| subdivision =
*''[[Aechmophorus]]''
*''[[Podiceps]]''
*''[[Podilymbus]]''
*''[[Poliocephalus]]''
*''[[Rollandia (bird)|Rollandia]]''
*''[[Tachybaptus]]''
}}
'''Grebes''' are members of the '''Podicipediformes''' [[order (biology)|order]], a widely distributed order of freshwater diving [[Aves|bird]]s, some of which visit the sea when [[Bird migration|migrating]] and in winter. This order contains only a single [[family (biology)|family]], the '''Podicipedidae''', containing 22 [[species]] in 6 extant [[genus|genera]].
==Description==
Grebes are small to medium-large in size, have lobed toes, and are excellent swimmers and divers. However, they have their feet placed far back on the body, can run for a short distance, but often fall over.
Grebes have narrow wings, and some species are reluctant to fly; indeed, two [[South America]]n species are completely flightless. They respond to danger by diving rather than flying, and are in any case much less wary than [[duck]]s. 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).
However, the [[North America]]n and [[Eurasia]]n 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; 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 [[Evolution|evolved]] independently in the extinct [[Cretaceous]]-age [[Hesperornithiformes]], which are totally unrelated birds.
Grebes have unusual [[feather|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.
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 ''[[Podiceps]]'' genus, are often striped and retain some of their juvenile plumage even after reaching full size.
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 [[parasite]]s.
==Taxonomy==
The grebes are a radically distinct group of birds as regards their [[anatomy]]. Accordingly, they were at first believed to be related to the [[loon]]s, which are also foot-propelled diving birds. However, as recently as the 1930s (Stolpe 1935), this was determined to be an example of [[convergent evolution]] by the strong [[selection|selective]] forces encountered by unrelated birds sharing the same lifestyle at different times and in different habitat.
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]] (Cracraft, 1982). 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) and [[DNA sequence|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.
However, because of the recent improvements in sequencing technology, the question of the taxonomy of grebes has finally been resolved. A recent [[phylogenomics|phylogenomic]] study of the deep evolutionary relationships in birds shows definitively that grebes are most closely related to [[flamingos]]. By collecting a large DNA sequence data bank, scientists was able to address the phylogenomic relationships from 171 species which represented all major [[passerine]] clades, all but three nonpasserine families, and two crocodile [[outgroups]] . All major extant groups were represented in this study, where they examined ~32 kilobases of aligned nuclear DNA sequences from 19 different [[loci]] for the 169 bird species plus the two crocodile outgroups (Hackett et al. 2008). This study validates earlier proposals of this relationship that have generally been ignored, even though careful analyses of morphological characters (i.e., excluding known [[homoplasies]]) and molecular data had suggested that this relationship was the most promising hypothesis to date. In addition, it was shown that the ''[[Anaticola]]'' [[bird louse|bird lice]] shared by flamingos and [[waterfowl]], which were at one time used to support a close relationship between the latter two (which is now deemed utterly incorrect), are actually closely related to the grebes' ''[[Aquanirmus]]'' lice and almost certainly switched hosts ''from'' flamingos ''to'' waterfowl, not the other way around as it was usually assumed (Johnson ''et al.'' 2006).
<!-- Auk123:1183; ZoolJLinnSoc140:157; -->
===Genera and species===
* Genus ''[[Tachybaptus]]''
**[[Little Grebe]], ''Tachybaptus ruficollis''
**[[Australasian Grebe]] ''Tachybaptus novaehollandiae''
**[[Madagascar Grebe]], ''Tachybaptus pelzelnii ''
**[[Alaotra Grebe]], ''Tachybaptus rufolavatus'' — probably [[extinct]] (late 1980s)
**[[Least Grebe]], ''Tachybaptus dominicus
* Genus ''[[Podilymbus]]''<!-- needs genus page; redirects to Pied-billed at present! -->
** [[Pied-billed Grebe]], ''Podilymbus podiceps''
** [[Atitlán Grebe]], ''Podilymbus gigas'' — extinct (1989)
* Genus ''[[Rollandia (bird)|Rollandia]]''
** [[White-tufted Grebe]], ''Rollandia rolland ''
** [[Titicaca Flightless Grebe]], ''Rollandia microptera ''
* Genus ''[[Poliocephalus]]''
** [[Hoary-headed Grebe]], ''Poliocephalus poliocephalus ''
** [[New Zealand Dabchick]], ''Poliocephalus rufopectus ''
* Genus ''[[Podiceps]]''
**[[Red-necked Grebe]], ''Podiceps grisegena''
**[[Great Crested Grebe]], ''Podiceps cristatus''
**[[Slavonian Grebe]] or Horned Grebe, ''Podiceps auritus''
**[[Black-necked Grebe]] or Eared Grebe, ''Podiceps nigricollis''
**[[Colombian Grebe]], ''Podiceps andinus'' — extinct (1977)
**[[Great Grebe]], ''Podiceps major ''
**[[Silvery Grebe]], ''Podiceps occipitalis ''
**[[Junin Flightless Grebe]], ''Podiceps taczanowskii ''
**[[Hooded Grebe]], ''Podiceps gallardoi''
* Genus ''[[Aechmophorus]]''
**[[Western Grebe]], ''Aechmophorus occidentalis''
**[[Clark's Grebe]], ''Aechmophorus clarkii''
===Fossil grebes===
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.{{Fact|date=May 2008}} The enigmatic waterbird [[genus]] ''[[Juncitarsus]]'', however, may be close to a common ancestor of flamingos and grebes.
The [[Late Cretaceous]] ([[Berriasian]], around 143 [[mya (unit)|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 (phylogenetics)|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 diagnositc and may not be of a bird at all (Benton ''et al.'', 1997).
''[[Telmatornis]]'' from the [[Navesink Formation]] - also Late Cretaceous - is traditionally allied with the [[Charadriiformes]] and/or [[Gruiformes]].<!-- AmMusNovit2388; Auk47:186; Auk84:260; Auk89:36; Condor33:107 -->. However, a [[cladistic]] analysis of the forelimb skeleton (Varricchio 2002) found it highly similar to the [[Great Crested Grebe]] and unlike the [[Painted Buttonquail]] (now known to be a [[basal (evolution)|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 (unit)|mya]]. While there are a few prehistoric genera that are now completely extinct, ''[[Thiornis]]'' (Late Miocene -? Early Pliocene of Libros, Spain)<!-- AnnPaleontol86:129; BullAMNH151:1--> and ''[[Pliolymbus]]'' (Late Pliocene of WC USA - Early? Pleistocene of Chapala, Mexico)<!-- Condor69:277; ContribMusPaleontolUnivMich24:205 --> date from a time when most if not all extant genera were already present. Only the Early Miocene ''[[Miobaptus]]'' from [[Czechoslovakia]]<!-- AnnPaleontol86:129 --> might be somewhat closer to the ancestral grebes, but more probably belongs to an extinct lineage. Indeed, ''Miobaptus'' is rivalled or even exceeded in age by a species of the modern genus ''[[Podiceps]]''.
A few more recent grebe fossils could not be assigned to modern or prehistoric genera as of now:
*Podicipedidae gen. et sp. indet. (San Diego Late Pliocene of California) - formerly included in ''Podiceps parvus'' (Murray 1967)
*Podicipedidae gen. et sp. indet. UMMP 49592, 52261, 51848, 52276, KUVP 4484 (Late Pliocene of WC USA: Jehl 1967)
*Podicipedidae gen. et sp. indet. (Glenns Ferry Late Pliocene/Early Pleistocene of Idaho, USA: Wetmore 1933, Jehl 1967)
== References ==
* {{aut|Benton, M. J.; Cook, E.; Grigorescu, D.; Popa, E. & Tallódi, E.}} (1997): Dinosaurs and other tetrapods in an Early Cretaceous bauxite-filled fissure, northwestern Romania. ''Palaeogeography, Palaeoclimatology, Palaeoecology'' '''130''': 275-292. <small>{{doi|10.1016/S0031-0182(96)00151-4}}</small> [http://palaeo.gly.bris.ac.uk/Benton/reprints/1997Romania.pdf PDF fulltext]
* {{aut|Cracraft, Joel}} (1982): Phylogenetic relationships and monophyly of loons, grebes, and hesperornithiform birds, with comments on the early history of birds. ''Systematic Zoology'' '''31''': 35-56. <small>{{DOI|10.2307/2413412}} </small> (HTML abstract)
* {{aut|Jehl, Joseph R. Jr.}} (1967): Pleistocene Birds from Fossil Lake, Oregon. ''[[Condor (journal)|Condor]]'' '''69'''(1): 24-27 [http://elibrary.unm.edu/sora/Condor/files/issues/v069n01/p0024-p0027.pdf PDF fulltext]
* {{aut|Johnso, Kevin P.; Kennedy, Martyn & McCracken, Kevin G.}} (2006): Reinterpreting the origins of flamingo lice: cospeciation or host-switching? ''[[Biology Letters|Biol. Lett.]]'' '''2''': 275–278. <small>{{DOI|10.1098/rsbl.2005.0427}}</small> [http://mercury.bio.uaf.edu/~kevin_mccracken/reprints/biol-lett-2-275.pdf PDF fulltext]
* {{aut|Hackett, S. J.; Kimball, R. T.; Reddy, S.; Bowie, R. C. K.; Braun, E. L.; Braun, M. J.; Chojnowski, J. L.; Cox, A.; Han, K.; Harshman, J.; Huddleston, C. J.; Marks, B. D.; Miglia, K. J.; Moore, W. S.; Sheldon, F. H.; Steadman, D. W.; Witt, C. C. & Tamaki, Y}} (2008):A Phylogenomic Study of Birds Reveals Their Evolutionary History. ''Science'' '''320''': 1763 - 1768. <small>{{doi: 10.1126/science.1157704}}</small> [http://www.sciencemag.org/cgi/reprint/320/5884/1763.pdf PDF fulltext]
* {{aut|Konter, André}} (2001): ''Grebes of our world: visiting all species on 5 continents''. Lynx Edicions, Barcelona. <small>ISBN 84-87334-33-4</small>
* {{aut|Murray, Bertram G. Jr.}} (1967): Grebes from the Late Pliocene of North America. ''[[Condor (journal)|Condor]]'' '''69'''(3): 277-288. [http://elibrary.unm.edu/sora/Condor/files/issues/v069n03/p0277-p0288.pdf PDF fulltext]
* {{aut|[[Malcolm Ogilvie|Ogilvie, Malcolm]] & Rose, Chris}} (2003): ''Grebes of the World''. Bruce Coleman Books, Uxbridge, England. <small>ISBN 1-872842-03-8</small>
* {{aut|[[Charles Sibley|Sibley, Charles Gald]] & Monroe, Burt L. Jr.}} (1990): ''Distribution and taxonomy of the birds of the world: A Study in Molecular Evolution''. Yale University Press, New Haven, CT. <small>ISBN 0-300-04969-2</small>
* {{aut|Stolpe, M.}} (1935): ''Colymbus, Hesperornis, Podiceps'':, ein Vergleich ihrer hinteren Extremität. ''[[Journal of Ornithology|J. Ornithol.]]'' '''80'''(1): 161-247. [Article in German] <small>{{DOI|10.1007/BF01908745}}</small>
* {{aut|Varricchio, David J.}} (2002): A new bird from the Upper Cretaceous Two Medicine Formation of Montana. ''Canadian Journal of Earth Sciences'' '''39'''(1): 19-26. [http://pubs.nrc-cnrc.gc.ca/cgi-bin/rp/rp2_abst_e?cjes_e01-057_39_ns_nf_cjes HTML abstract]
* {{aut|Wetmore, Alexander}} (1933): Pliocene bird remains from Idaho. ''Smithsonian Miscellaneous Collections'' '''87''': 1-12.
==External links==
*Internet Bird Collection: [http://ibc.hbw.com/ibc/phtml/familia.phtml?idFamilia=9 Grebe videos]
* [http://www.tolweb.org/Podicipedidae/26371 Tree of Life Grebes]
*openclipart.org: [http://openclipart.org/cchost/media/tags/aechmophorus Grebes clip-art]
* [http://digitalcommons.unl.edu/bioscidivingbirds/ ''Diving Birds of North America'', by Paul Johnsgard]
{{Footer Birds}}
[[Category:Podicipedidae]]
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