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Megapode
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{{Short description|Family of birds}} {{distinguish|text=[[Macropod (disambiguation){{!}}Macropod]], kangaroos and similar marsupials}} {{Automatic taxobox | image = Australian brushturkey (Alectura lathami) female Atherton.jpg | image_caption = female [[Australian brushturkey]] (''Alectura lathami'') | taxon = Megapodiidae | authority = [[René-Primevère Lesson|Lesson]], 1831 | subdivision_ranks = [[Genera]] | subdivision = *''[[Aepypodius]]'' *''[[Alectura]]'' *''[[Eulipoa]]'' *''[[Leipoa]]'' *''[[Macrocephalon]]'' *''[[Megapodius]]'' *''[[Talegalla]]'' }} [[File:Australian brushturkey (Alectura lathami) female head Atherton.jpg|thumb|right|Brushturkeys can often be found in parks or gardens.]] The '''megapodes''', also known as '''incubator birds''' or '''mound-builders''', are stocky, medium-large, [[chicken]]-like [[bird]]s with small heads and large feet in the family '''Megapodiidae'''. Their name literally means "large foot" and is a reference to the heavy legs and feet typical of these [[Terrestrial animal|terrestrial]] birds. All are [[Browsing (predation)|browsers]], and all except the [[malleefowl]] occupy [[Woodland|wooded]] habitats. Most are brown or black in color. Megapodes are [[superprecocial]], hatching from their eggs in the most mature condition of any bird. They hatch with open eyes, bodily coordination and strength, full wing feathers, and downy body feathers, and are able to run, pursue prey and, in some species, fly on the day they hatch.<ref name=Starck-Ricklefs-1998/> ==Etymology== From the Greek {{lang|grk|μέγας}} ({{transliteration|grk|mégas}} = ''great'') and {{lang|grk|πούς}}, ({{transliteration|grk|poús}} = ''foot'').<ref>{{cite EB1911 |wstitle=Megapode |volume=18 |page=75|first=Alfred |last=Newton}}</ref> ==Description== Megapodes are medium-sized to large terrestrial birds with large legs and feet with sharp claws. Megapodes are of three kinds: scrub fowl, brush turkeys, and mallee fowl or lowan. The largest members of the [[clade]] are the species of ''[[Alectura]]'' and ''[[Talegalla]]''. The smallest are the [[Micronesian scrubfowl]] (''Megapodius laperouse'') and the [[Moluccan scrubfowl]] (''Eulipoa wallacei''). They have small heads, short beaks, and rounded and large wings. Their flying abilities vary within the clade. They present the [[hallux#In other animals|hallux]] at the same level of the other toes just like the species of the clade [[Cracidae]]. The other [[Galliformes]] have their halluces raised above the level of the front toes.<ref name=delHoyo-Elliott-Sargatal-1994/> == Distribution and habitat== Megapodes are found in the broader [[Australasia]]n region, including islands in the western [[Pacific Ocean|Pacific]], [[Australia]], [[New Guinea]], and the islands of [[Indonesia]] east of the [[Wallace Line]], but also the [[Andaman Islands|Andaman]] and [[Nicobar Islands]] in the [[Bay of Bengal]]. The distribution of the family has contracted in the Pacific with the arrival of humans, and a number of island groups such as [[Fiji]], [[Tonga]], and [[New Caledonia]] have lost many or all of their species. [[Raoul Island]], a New Zealand territory and the main island of the [[Kermadec Islands]], may also have once had a species of megapode, based on settler accounts.<ref name=Steadman-2006/> ==Behaviour and ecology== [[Image:Brushturkeykansaszoo.jpg|thumb|right|250px|Australian brushturkey on its mound]] Megapodes are mainly solitary birds that do not incubate their eggs with their body heat as other birds do, but bury them. Their eggs are unusual in having a large yolk, making up 50–70% of the egg weight.<ref name=Starck-Sutter-2000/> The birds are best known for building massive nest mounds of decaying vegetation, which the male attends, adding or removing litter to regulate the internal heat while the eggs develop. However, some bury their eggs in other ways; there are burrow-nesters which use geothermal heat, and others which simply rely on the heat of the sun warming the sand. Some species vary their incubation strategy depending on the local environment.<ref name=Steadman-2006/> Although the [[Australian brushturkey]] was thought to exhibit [[temperature-dependent sex determination]], this was later proven false;<ref name=Göth-Booth-2005-03-22/> temperature does, however, affect embryo mortality and resulting offspring sex ratios. The nonsocial nature of their incubation raises questions as to how the hatchlings come to recognise other members of their species, which is due to [[imprinting (psychology)|imprinting]] in other members of the order [[Galliformes]]. Research suggests an instinctive visual recognition of specific movement patterns is made by the individual species of megapode.<ref name=Göth-Evans-2004/> [[Image:Malleefowl Pengo.svg|thumb|right|250px|This cross-section of a megapode mound shows a layer of sand (up to 1 m thick) used for insulation, an egg chamber, and a layer of rotting [[compost]]. The egg chamber is kept at a constant 33°C by opening and closing air vents in the insulation layer, while heat comes from the compost below.]] Megapode chicks do not have an [[egg tooth]]; they use their powerful claws to break out of the egg, and then tunnel their way up to the surface of the mound, lying on their backs and scratching at the sand and vegetable matter. Similar to other [[superprecocial]] birds, they hatch fully feathered and active, already able to fly and live independently from their parents.<ref name=Starck-Sutter-2000/> In megapodes superprecociality apparently evolved secondarily from brooding and at least loose parental care as more typical in Galliformes.<ref name=Harris-Birks-Leaché-2014/> Eggs previously assigned to ''[[Genyornis]]'' have been reassigned to giant megapode species. Some dietary and chronological data previously assigned to [[Dromornithidae|dromornithids]] may instead be assigned to the giant megapodes.<ref name=Worthy-2016-01-13/> Megapodes share some similarities to the extinct [[enantiornithes]] in terms of their superprecocial life cycle, though also several differences.{{efn| "These feather traces and the plumage in HPG-15-1 strongly suggest that members of the Enantiornithes were born fully fledged and capable of flight soon after hatching, somewhat resembling the super-precocial megapodes, the only group of neornithines in which neonates are similarly born fledged and capable of flight (Zhou and Zhang, 2004; Jones and Göth, 2008; Xing et al., 2017). Megapodes do not fly immediately, requiring nearly two days to dig themselves out of their mounds during which they preen off their feather sheaths and let their feathers dry (Jones and Göth, 2008). Similarly, hatchling enantiornithines would have had to wait until their feather sheaths were removed and their feathers dry before attempting flight. Although ecological and behavioural differences clearly exist between enantiornithines and megapodes (e.g., enantiornithines were arboreal and not mound-nesters), megapodes represent the precocial extreme in extant neornithines and thus the closest analogue for enantiornithine development, for which all evidence indicates a form of extreme precociality (Elzanowski, 1981; Zhou and Zhang, 2004; Xing et al., 2017)."<ref name=OConnor-Falk-etal-2020-01-03/> }} ==Species== The more than 20 living species are placed in seven genera. Although the evolutionary relationships between the Megapodiidae are especially uncertain, the morphological groups are clear:<ref name=Birks-Edwards-2002/> ===Phylogeny=== {{Cladogram |caption=Phylogeny of the Megapodiidae based a study by Chen, Hosner, Dittmann, ''et al''. (2021).<ref name=Chen-Hosner-etal-2021/> The numbers of species are from the [[International Ornithologists' Union|IOU]] list by [[Frank Gill (ornithologist)|Gill]], Donsker, & [[Pamela Rasmussen|Rasmussen]].<ref name=Gill-Donsker-Rasmussen-2021-07/> |align=centre |cladogram={{Clade| style=font-size:100%;line-height:80%;width:550px; |1={{clade |label1=Megapodiidae |1={{clade |1={{clade |1={{clade |1=''[[Macrocephalon]]'' – maleo |2={{clade |1=''[[Eulipoa]]'' – Moluccan megapode |2=''[[Megapodius]]'' – scrubfowl (12 species) }} }} |2={{clade |1=''[[Leipoa]]'' – malleefowl |2={{clade |1=[[Talegalla]] – brushturkeys (3 species) |2={{clade |1=''[[Alectura]]'' – Australian brushturkey |2=''[[Aepypodius]]'' – brushturkeys (2 species) }} }} }} }} }} }} }} }} ===Taxonomy=== * Genus †''[[Mwalau]]'' <small>Worthy et al. 2015</small> ** †''[[Mwalau walterlinii]]'' <small>Worthy et al. 2015</small> (Vanuatu) * Genus †''[[Ngawupodius]]'' <small>Boles & Ivison 1999</small> ** †''[[Ngawupodius minya]]'' <small>Boles & Ivison 1999</small> * '''[[Scrubfowl|Scrubfowl group]]''' ** Genus: ''Macrocephalon'' *** [[Maleo]], ''Macrocephalon maleo'' ** Genus: ''[[Eulipoa]]'' (sometimes included in ''Megapodius'') *** [[Moluccan megapode]], ''Eulipoa wallacei''. ** Genus: ''[[Megapodius]]'' *** [[Tongan megapode]], ''Megapodius pritchardii'' *** [[Micronesian megapode]], ''Megapodius laperouse'' **** [[Marianas Island megapode]], ''Megapodius laperouse laperouse'' **** [[Palau Island megapode]], ''Megapodius laperouse senex'' *** [[Nicobar megapode]], ''Megapodius nicobariensis'' *** [[Philippine megapode]], ''Megapodius cumingii'' *** [[Sula megapode]], ''Megapodius bernsteinii'' *** [[Tanimbar megapode]], ''Megapodius tenimberensis'' *** [[Dusky megapode]], ''Megapodius freycinet'' **** [[Forsten's megapode]], ''Megapodius (freycinet) forstenii'' *** [[Biak scrubfowl]], ''Megapodius geelvinkianus'' *** [[Melanesian megapode]], ''Megapodius eremita'' *** [[Vanuatu megapode]], ''Megapodius layardi'' *** [[New Guinea scrubfowl]], ''Megapodius decollatus'' *** [[Orange-footed scrubfowl]], ''Megapodius reinwardt'' *** †[[Pile-builder scrubfowl]], ''Megapodius molistructor'' <small>Balouet & Olson 1989</small> *** †[[Viti Levu scrubfowl]], ''Megapodius amissus'' <small>Worthy 2000</small> *** †[[Consumed scrubfowl]], ''Megapodius alimentum'' <small>Steatman 1989a</small> *** †''M. andamanensis'' <small>Walter 1980 nomen dubium</small> [oospecies] *** †''M. burnabyi'' <small>Gray 1861 nomen dubium</small> [oospecies] *** †Raoul Island scrubfowl, ''M. sp.'' *** †'Eua scrubfowl (small-footed megapode), ''M. sp.'' *** †Lifuka scrubfowl, ''M. sp.'' *** †Stout Tongan megapode, ''M. sp.'' *** †Large Solomon Islands megapode, ''M. sp.'' *** †New Caledonia megapode, ''M. sp.'' *** †Loyalty megapode, ''M. sp.'' *** †New Ireland scrubfowl (large Bismarck's megapode), ''M. sp.'' * '''[[Malleefowl]], group''' ** Genus: ''Leipoa'' *** [[Malleefowl]], ''Leipoa ocellata'' * '''[[Brushturkey|Brushturkey group]]''' ** Genus: ''Alectura'' *** [[Australian brushturkey]], ''Alectura lathami'' ** Genus: ''[[Aepypodius]]'' *** [[Wattled brushturkey]], ''Aepypodius arfakianus'' *** [[Waigeo brushturkey]], ''Aepypodius bruijnii'' ** Genus: ''[[Talegalla]]'' *** [[Red-billed brushturkey]], ''Talegalla cuvieri'' *** [[Black-billed brushturkey]], ''Talegalla fuscirostris'' *** [[Collared brushturkey]], ''Talegalla jobiensis'' ** Genus: †''[[Progura]]'' *** ''Progura gallinacea'' – Queensland, Pleistocene *** ''Progura campestris'' – South Australia, Pleistocene ** Genus: †''[[Latagallina]]'' *** ''Latagallina naracoortensis'' formerly ''Progura naracoortensis'' – New South Wales, South Australia, Pleistocene *** ''Latagallina olsoni'' – South Australia, Pleistocene * ''[[Incertae sedis]]'' ** Genus: †''[[Garrdimalga]]'' *** ''Garrdimalga mcnamarai'' – South Australia, Pleistocene ==Human uses== In their native [[Oceania]], indigenous peoples protect their nesting sites, as their eggs are considered to be delicacies.<ref>{{cite book |title=Microlivestock: Little-known small animals with a promising economic future |year=1991 |publisher=National Academies Press |isbn=978-0309042956 |pages=170–171 |doi=10.17226/1831 |url=https://nap.nationalacademies.org/read/1831/chapter/17 |access-date=5 April 2022}}</ref> Their eggs are about twice the size of chicken eggs and the yolks are roughly four times as massive.<ref>{{cite web |last1=Cocker |first1=Mark |title=The Lava Birds |url=https://www.visitsolomons.com.sb/wp-content/uploads/2019/06/BBC-Wildlife-megapodes.pdf |website=Tourism Solomons |access-date=5 April 2022}}</ref> ==See also== * [[List of recently extinct birds]] * [[Late Quaternary prehistoric birds]] * [[List of fossil bird genera]] == Footnotes == {{notelist}} == References == {{reflist|22em|refs= <ref name=Birks-Edwards-2002> {{cite journal |author1=Birks, S.M. |author2=Edwards, S.V. |name-list-style=amp |year=2002 |title=A phylogeny of the megapodes (Aves: Megapodiidae) based on nuclear and mitochondrial DNA sequences |journal=Molecular Phylogenetics and Evolution |volume=23 |issue=3 |pages=408–421 |doi=10.1016/s1055-7903(02)00002-7 |pmid=12099795 |bibcode=2002MolPE..23..408B }} </ref> <ref name=Chen-Hosner-etal-2021> {{cite journal |last1=Chen |first1=D. |last2=Hosner |first2=P.A. |last3=Dittmann |first3=D.L. |last4=O’Neill |first4=J.P. |last5=Birks |first5=S.M. |last6=Braun |first6=E.L. |last7=Kimball |first7=R.T. |date=2021 |title=Divergence time estimation of Galliformes based on the best gene shopping scheme of ultraconserved elements |journal=BMC Ecology and Evolution |volume=21 |issue=1 |page=209 |doi=10.1186/s12862-021-01935-1 |doi-access=free |pmid=34809586 |pmc=8609756 }} </ref> <ref name=delHoyo-Elliott-Sargatal-1994> {{cite book |first1=J. |last1=del Hoyo |first2=A. |last2=Elliott |first3=J. |last3=Sargatal |year=1994 |title=Handbook of the Birds of the World |volume=2: New World Vultures to Guineafowl |publisher=Lynx Edicions |isbn=978-84-87334-15-3 |url-access=registration |url=https://archive.org/details/handbookofbirdso0001unse }} </ref> <ref name=Gill-Donsker-Rasmussen-2021-07> {{cite web |editor1-last=Gill |editor1-first=Frank |editor1-link=Frank Gill (ornithologist) |editor2-last=Donsker |editor2-first=David |editor3-last=Rasmussen |editor3-first=Pamela |editor3-link=Pamela Rasmussen |date=July 2021 |title=Pheasants, partridges, francolins |series=IOC World Bird List |edition=Version 11.2 |publisher=[[International Ornithologists' Union]] |url=https://www.worldbirdnames.org/bow/pheasants/ |access-date=23 November 2021 }} </ref> <ref name=Göth-Evans-2004> {{cite journal |author1=Göth, A. |author2=Evans, C.S. |name-list-style=amp |year=2004 |title=Social responses without early experience: Australian brush-turkey chicks use specific visual cues to aggregate with conspecifics |journal=Journal of Experimental Biology |volume=207 |issue=13 |pages=2199–2208 |doi=10.1242/jeb.01008 |doi-access=free |pmid=15159424 |bibcode=2004JExpB.207.2199G }} </ref> <ref name=Göth-Booth-2005-03-22> {{cite journal |last1=Göth |first1=Ann |last2=Booth |first2=David T. |name-list-style=amp |date=22 March 2005 |title=Temperature-dependent sex ratio in a bird |journal=Biology Letters |volume=1 |issue=1 |pages=31–33 |doi=10.1098/rsbl.2004.0247 |pmid=17148121 |pmc=1629050 }} </ref> <ref name=Harris-Birks-Leaché-2014> {{cite journal |last1 = Harris |first1 = R.B. |last2 = Birks |first2 = S.M. |last3 = Leaché |first3 = A.D. |year = 2014 |title = Incubator birds: Biogeographical origins and evolution of underground nesting in megapodes (Galliformes: Megapodiidae) |journal = Journal of Biogeography |volume = 41 |issue = 11 |pages = 2045–2056 |doi = 10.1111/jbi.12357 |bibcode = 2014JBiog..41.2045H |s2cid = 53532720 }} </ref> <ref name=OConnor-Falk-etal-2020-01-03> {{cite journal |first1=Jingmai |last1=O'Connor |first2=Amanda |last2=Falk |first3=Min |last3=Wang |first4=Zheng |last4=Xiao-Ting |date=3 January 2020 |title=First report of immature feathers in juvenile enantiornithines from the Early Cretaceous Jehol avifauna |doi=10.19615/j.cnki.1000-3118.190823 |series=Project: Vertebrata PalAsiatica |url=https://www.researchgate.net/publication/338355119 |website=ResearchGate }} </ref> <ref name=Starck-Ricklefs-1998> {{cite book |author1=Starck, J.M. |author2=Ricklefs, R.E. |name-list-style=amp |year=1998 |title=Avian Growth and Development: Evolution within the altricial precocial spectrum |place=New York, NY |publisher=[[Oxford University Press]] |isbn=978-0-19-510608-4 }} </ref> <ref name=Starck-Sutter-2000> {{cite journal |author1=Starck, J.M. |author2=Sutter E. |name-list-style=amp |year=2000 |title=Patterns of growth and heterochrony in moundbuilders (MEgapodiidae) and fowl (Phasianidae) |journal=Journal of Avian Biology |volume=31 |issue=4 |pages=527–547 |doi=10.1034/j.1600-048x.2000.310413.x }} </ref> <ref name=Steadman-2006> {{cite book |last=Steadman |first=D. |year=2006 |title=Extinction and Biogeography in Tropical Pacific Birds |publisher=University of Chicago Press |isbn=978-0-226-77142-7 }} </ref> <ref name=Worthy-2016-01-13> {{cite web |first=Trevor H. |last=Worthy |date=13 January 2016 |title=A case of mistaken identity for Australia's extinct big bird |website=theconversation.com |url=http://theconversation.com/a-case-of-mistaken-identity-for-australias-extinct-big-bird-52856 }} </ref> }} <!-- end "refs=" --> == External links == {{Commons category|Megapodiidae}} * {{cite AV media |title=Mound-builders |medium=videos, photos, & sounds |website=Internet Bird Collection (ibc.lynxeds.com) |url=http://ibc.lynxeds.com/family/megapodes-megapodiidae}} * {{cite AV media |title=Nest mound of ''M. tenimberensis'' |medium=photograph |publisher=Oriental Bird Club |via=Internet Archive (archive.org) |url=http://orientalbirdimages.org/photographers.php?p=3&action=birderimages&Bird_Image_ID=26455&Birder_ID=693&PHPSESSID=1ae |url-status=dead <!-- presumed --> |archive-url=https://web.archive.org/web/20110727154553/http://orientalbirdimages.org/photographers.php?p=3&action=birderimages&Bird_Image_ID=26455&Birder_ID=693&PHPSESSID=1ae |archive-date=2011-07-27}} {{Birds|all}} {{Taxonbar|from=Q213536}} {{Authority control}} [[Category:Megapodiidae|*]] [[Category:Taxa named by René Lesson]]
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