Gen Anderson and Chris Holmes Galliformes are robust, ground‐dwelling birds with stubby, rounded wings adapted for short flights versus long distance dispersal. They are found nearly globally in a diverse variety of terrestrial habitats such as the ptarmigan (Lagopus sp.) living within the Arctic Circle, the cloud forest‐residing horned guan (Oreophasis derbianus), and the desert‐loving vulturine guineafowl (Acryllium vulturinum). Galliformes generally have a rotund body shape with stout legs and an anisodactyl toe arrangement ideal for scratching around in substrate, and are sexually dimorphic. Many species roost arboreally at night for safety. They typically are opportunistic omnivores, foraging on the ground for invertebrates, seeds, fallen fruits, tender leaves, and tubers. They serve an important role in the ecosystem as seed dispersers. Many species are highly valued economically for hunting and as a food source. Aside from humans, Galliformes are heavily preyed upon by most terrestrial carnivores. Multiple species within the order Galliformes have been domesticated for cockfighting, ornamental display and to supply meat, eggs, and feathers. Examples include helmeted guineafowl, Indian peafowl, wild turkey, bobwhite quail and the domestic chicken [1]. Recent genome sequencing analysis points to the red junglefowl, Gallus g. spadiceus, of China, Thailand and Myanmar as the primary source of origin for domestic chicken populations. The species split occurred roughly 9500 (±3300) years ago, with the domestication process progressing during the Holocene period. Wild populations of other Gallus species and subspecies have hybridized incrementally with free‐ranging domestic birds [2]. Hundreds of chicken breeds are recognized worldwide. They are often used in zoological parks as animal ambassadors, for natural pest control, and incubation and rearing of exotic Galliformes when appropriate [1]. Hybridization across species and color morphs within species of the Phasianidae family are abundant in private aviculture. Many were accidental crossings, but some were encouraged to increase ornamental appeal or economic profit through meat and egg production. Crossings of golden and Lady Amherst pheasants (Chrysolophus pictus x amherstiae) are far too common and unfortunately produce fertile offspring. Responsible breeders attempt to locate wild‐type birds, but pure species of either of these are few and far between. Golden pheasants have historically also been crossed with silver (Lophura nycthemera) and ring‐necked pheasants. Tragopan hens of differing species look nearly identical and have resulted in the production of hybrids by many a well‐intentioned aviculturist. Vietnam pheasants (Lophura edwardsi) were historically hybridized with L. nycthemera and Swinhoe’s pheasant (Lophura swinhoii). Indian and green peafowl crosses (Pavo cristatus x muticus) produce fertile hybrids referred to as Spalding peafowl. Domestic chickens, guineafowl, and other pheasant species are unfortunately still intentionally crossbred to this day [3]. The taxonomic and phylogenetic relationships within the Galliformes order have been studied by numerous researchers, but remain debated and ever evolving. Referenced publications for this chapter analyze data from mitochondrial and nuclear DNA, known fossil calibrations, region of origin, and/or morphological characteristics. Dissimilar chronological results differ due to sample set variations, fossil placement, amount of data used, taxonomic sample sizes, mitochondrial vs nuclear data, etc. [4]. The ancestral Galloanserae (fowl clade) is universally accepted to have split into Galliformes and Anseriformes during the Cretaceous period roughly 76 million years ago [9]. The Cretaceous‐ Paleogene (K‐Pg) mass extinction 66 million years ago devastated >75% of the planet’s flora and fauna, including dinosaurs, arboreal vertebrates, and forests. Few avian species survived but among them were ground‐dwelling birds, including Galloanserae and Palaeognathae (ratites), that went through massive adaptive radiation in the Paleogene period, resulting in the modern birds found today [5]. Jiang et al. estimated the family divergence of modern Galliformes to have occurred over 16–28 myr [6]. Cornell Lab of Ornithology’s Birds of the World (BOW) was selected to use as the most up‐to‐date taxonomic source. This online resource was recently developed using a combination of The Handbook of the Birds of the World Alive (HBW Alive), Birds of North America, and Neotropical Birds. The following editorial note is stated: “Due to the merger of content from various sources, especially related to differing taxonomies, there are cases where the written text does not perfectly match up with the current taxonomy, or in the worst cases the two may contradict each other” [7]. Every effort has been made to use the most current accepted taxonomy here for reference. Differing taxonomies result from lumping species or subspecies into larger groups or splitting them into multiple small groups. The living order Galliformes consists of two suborders, five families and 295 total species (Figure 3.1). The suborder Cracii is further divided into Megapodiidae (the megapodes, 22 species) and Cracidae (the chachalacas, guans, and curassows, 54 species). The suborder Phasiani is divided into Odontophoridae (the New World quails, 33 species), Numididae (the guineafowl, six species), and Phasianidae (pheasants, partridges, turkeys, and grouse, 180 species) [8]. This unusual family of extant Gallifomes contains 22 species within seven genera (Table 3.1). Morphologically large, stout birds from Australasian primarily inhabiting lowland tropical forests with the exception of the arid‐adapted malleefowl. Numerous species are island endemics. Their reproductive techniques are unique in Aves, constructing nest mounds or burrows that depend on heat produced by the breakdown of organic material, geothermal sources or solar power for egg incubation. Lacking an eggtooth, the hatchlings burst out of their shells using powerfully kicking feet. Young are superprecocial, requiring no parental care, and fly away shortly after hatch [7]. Megapodiidae are estimated to have diverged during the late Cretaceous, around 66.8 million years ago (mya) [9] before increasing diversification around the Eocene–Oligocene period transition 33.9 mya [4]. They are a sister clade to Cracidae [6]. More species have gone extinct than exist today. Judging by fossil records, known extinctions and species sympatry preference, 45–55 species of Megapodes are presumed to have existed historically prior to human impact caused by colonization of Oceania during the past several millennia [10]. Per the Species360 Zoological Information Management System (ZIMS), few megapode species are managed in aviculture and none in genetically sustainable population sizes. The maleo (Macrocephalon maleo) (Figure 3.2) and Australian brush turkey (Alectura lathami) represent most of the birds in aviculture. Malleefowl (Leipoa ocellata) and a single Niuafoou scrub fowl (Megapodius pritchardii) are also maintained in insignificant numbers [11]. Table 3.1 Megapodiidae family with genera. Residing only in the New World, the chachalacas, guans, and curassows include 54 species in 11 genera (Table 3.2, Figure 3.3). Shared characteristics for this family include larger, heavy‐bodied birds typically with elaborate head adornments in the form of crests, horns, wattles or casques. Though their wings are short and round, similar to other Galliformes, the tails of cracids are long and they are capable of strong flight. Prominent vocalizations include booming, hums, rattles, clucks, yelps, and whistles [7]. Table 3.2 Cracidae family with genera. Cracidae are one of the earliest diverging families along with sister clade Megapodiidae, which together are sister clades to Numididae [6, 9]. Cracidae diverged either immediately prior to the Cretaceous–Paleogene boundary or in the early Paleogene 50–60 mya. All remaining Galliformes divergences occurred during the Paleogene period [9]. Early fossil records pointed to origins in North America when the region was tropical. Expansion into Central and South America occurred as ancient climates changed [12]. On the other hand, using a combination of mitochondrial and nuclear data with fossil calibrations, ancestral origins point to South America or dispersal from Africa when those continents were much closer than at present [9]. Cracids are fairly common in aviculture, though none of the species can be found in robust population sizes. In North America, most Crax sp. are maintained, but primarily only one guan species, the crested guan (Penelope purpurascens). All piping guan species (Pipile sp.) are extremely common in aviculture. Chaco chacalacas (Ortalis canicollis) are in greater numbers in Europe, but also are managed in the United States in very small numbers along with the plain chachalaca (Ortalis vetula). The distinctive horned guan (O. derbianus) and blue‐billed curassows (Crax alberti) are only found within American Association of Zoos and Aquariums (AZA)‐accredited zoological parks [11]. Composed of four genera and six species, this is one of the smallest Galliformes family groups (Table 3.3, Figure 3.4). All species reside in coastal and sub‐Saharan Africa. Shared characteristics within Numididae typically include dark plumage with boldly distinctive spots and/or stripes. These taxa display brightly colored heads and facial wattles. All species are sexually monomorphic. Guineafowl were historically classified in Phasianidae, but were broken out by molecular phylogenetic studies [7]. Table 3.3 Numidae family with genera.
3
Galliformes: Species, Varieties, and Behavior
3.1 Introduction
3.2 Domestication and Hybridization
3.3 Systematics
3.4 Megapodiidae – Megapodes
Family
Genus
Common name
Latin name
Number of subspecies
Described by
Date
Megapodiidae
Alectura
Australian brush turkey
Alectura lathami
2
J.E. Gray
1831
Aepypodius
Wattled brush turkey
Aepypodius arfakianus
2
Salvadori
1877
Waigeo brush turkey
Aepypodius bruijnii
Monotypic
Oustalet
1880
Talegalla
Red‐billed brush turkey
Talegalla cuvieri
2
Lesson
1828
Yellow‐billed brush turkey
Talegalla fuscirostris
4
Salvadori
1877
Red‐legged brush turkey
Talegalla jobiensis
2
A.B. Meyer
1874
Leipoa
Malleefowl
Leipoa ocellata
Monotypic
Gould
1840
Macrocephalon
Maleo
Macrocephalon maleo
Monotypic
S. Müller
1846
Eulipoa
Moluccan scrubfowl
Eulipoa wallacei
Monotypic
G.R. Gray
1861
Megapodius
Niuafoou scrubfowl
Megapodius pritchardii
Monotypic
G.R. Gray
1864
Micronesian scrubfowl
Megapodius laperouse
2
Gaimard
1823
Nicobar scrubfowl
Megapodius nicobariensis
2
Blyth
1846
Tabon scrubfowl
Megapodius cumingii
7
Dillwyn
1853
Sula scrubfowl
Megapodius bernsteinii
Monotypic
Schlegal
1866
Tanimbar scrubfowl
Megapodius tenimberensis
Monotypic
P.L. Sclater
1883
Dusky scrubfowl
Megapodius freycinet
3
Gaimard
1823
Biak scrubfowl
Megapodius geelvinkianus
Monotypic
A.B. Meyer
1874
Forsten’s scrubfowl
Megapodius forsteni
2
G.R. Gray
1847
Melanesian scrubfowl
Megapodius eremita
Monotypic
Hartlaub
1866
Vanuatu scrubfowl
Megapodius layardi
Monotypic
Tristram
1879
New Guinea scrubfowl
Megapodius decollatus
Monotypic
Oustalet
1878
Orange‐footed scrubfowl
Megapodius reinwardt
5
Dumont de Sainte Croix
1823
3.5 Cracidae – Guans, Chachalacas, Curassows
Family
Genus
Common name
Latin name
Number of subspecies
Described by
Date
Cracidae
Ortalis
Plain chachalaca
Ortalis vetula
4
Wagler
1830
Gray‐headed chachalaca
Ortalis cinereiceps
Monotypic
G.R. Gray
1867
Chestnut‐winged chachalaca
Ortalis garrula
Monotypic
Humboldt
1805
Rufous‐vented chachalaca
Ortalis ruficauda
2
Jardine
1847
Rufous‐headed chachalaca
Ortalis erythroptera
Monotypic
P.L. Sclater & Salvin
1870
Rufous‐bellied chachalaca
Ortalis wagleri
Monotypic
G.R. Gray
1867
West Mexican chachalaca
Ortalis poliocephala
Monotypic
Wagler
1830
Chaco chachalaca
Ortalis canicollis
2
Wagler
1830
White‐bellied chachalaca
Ortalis leucogastra
Monotypic
Gould
1843
Colombian chachalaca
Ortalis columbiana
Monotypic
Hellmayr
1906
Speckled chachalaca
Ortalis guttata
3
Spix
1825
East Brazilian chachalaca
Ortalis araucuan
Monotypic
Spix
1825
Scaled chachalaca
Ortalis squamata
Monotypic
Lesson
1829
Variable chachalaca
Ortalis motmot
2
Linnaeus
1766
Buff‐browed chachalaca
Ortalis superciliaris
Monotypic
G.R. Gray
1867
Penelope
Band‐tailed guan
Penelope argyrotis
3
Bonaparte
1856
Bearded guan
Penelope barbata
Monotypic
Chapman
1921
Baudo guan
Penelope ortoni
Monotypic
Salvin
1874
Andean guan
Penelope montagnii
5
Bonaparte
1856
Marail guan
Penelope marail
2
Statius Müller
1776
Rusty‐margined guan
Penelope superciliaris
3
Temminck
1815
Red‐faced guan
Penelope dabbenei
Monotypic
Hellmayr & Conover
1942
Spix’s guan
Penelope jacquacu
4
Spix
1825
Crested guan
Penelope purpurascens
3
Wagler
1830
Cauca guan
Penelope perspicax
Monotypic
Bangs
1911
White‐winged guan
Penelope albipennis
Monotypic
Taczanowski
1878
Dusky‐legged guan
Penelope obscura
3
Temminck
1815
White‐crested guan
Penelope pileata
Monotypic
Wagler
1830
Chestnut‐bellied guan
Penelope ochrogaster
Monotypic
Pelzeln
1870
White‐browed guan
Penelope jacucaca
Monotypic
Spix
1825
Pipile
Trinidad piping‐guan
Pipile pipile
Monotypic
Jacquin
1784
Blue‐throated piping‐guan
Pipile cumanensis
2
Jacquin
1784
Red‐throated piping‐guan
Pipile cujubi
2
Pelzeln
1858
Black‐fronted piping‐guan
Pipile jacutinga
Monotypic
Spix
1825
Aburria
Wattled guan
Aburria aburri
Monotypic
Lesson
1828
Chamaepetes
Black guan
Chamaepetes unicolor
Monotypic
Salvin
1867
Sickle‐winged guan
Chamaepetes goudotii
5
Lesson
1828
Penelopina
Highland guan
Penelopina nigra
Monotypic
Fraser
1852
Oreophasis
Horned guan
Oreophasis derbianus
Monotypic
G.R. Gray
1844
Nothocrax
Nocturnal curassow
Nothocrax urumutum
Monotypic
Spix
1825
Mitu
Crestless curassow
Mitu tomentosa
Monotypic
Spix
1825
Salvin’s curassow
Mitu salvini
Monotypic
Reinhardt
1879
Razor‐billed curassow
Mitu tuberosa
Monotypic
Spix
1825
Alagoas curassow
Mitu mitu
Monotypic
Linnaeus
1766
Pauxi
Helmeted curassow
Pauxi pauxi
2
Linnaeus
1766
Sira curassow
Pauxi koepckeae
Monotypic
Weske & Terborgh
1971
Horned curassow
Pauxi unicornis
Monotypic
J. Bond & Meyer de Schauensee
1939
Crax
Great curassow
Crax rubra
2
Linnaeus
1758
Blue‐billed curassow
Crax alberti
Monotypic
Fraser
1850
Yellow‐knobbed curassow
Crax daubentoni
Monotypic
G.R. Gray
1867
Black curassow
Crax alector
2
Linnaeus
1766
Wattled curassow
Crax globulosa
Monotypic
Spix
1825
Bare‐faced curassow
Crax fasciolata
3
Spix
1825
Red‐billed curassow
Crax blumenbachii
Monotypic
Spix
1825
3.6 Numididae – Guineafowl
Family
Genus
Common name
Latin name
Number of subspecies
Described by
Date
Numididae
Numida
Helemeted guineafowl
Numida meleagris
9
Linnaeus
1758
Agelastes
White‐breasted guineafowl
Agelastes meleagrides
Monotypic
Bonaparte
1850
Black guineafowl
Agelastes niger
Monotypic
Cassin
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