3Galliformes: Species, Varieties, and Behavior
Gen Anderson and Chris Holmes
3.1 Introduction
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.
3.2 Domestication and Hybridization
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].
3.3 Systematics
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].

Figure 3.1 Galliformes cladogram displaying the relationships among families of the Galliformes. The taxonomic and phylogenetic relationships within this order remain debated and ever evolving.
3.4 Megapodiidae – Megapodes
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.
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 |

Figure 3.2 Maleo (Macrocephalon maleo).
Source: Riza Marlon/Shutterstock.
3.5 Cracidae – Guans, Chachalacas, Curassows
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.
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 |

Figure 3.3 Cracidae. (a) Wattled curassow (Crax globulosa) hen.
Source: Carolyn Atherton, Audubon Zoo. (b) Horned guan (Oreophasis derbianus).
Source: JoEllen Toler, St Louis Zoo. (c) Plain chachalaca (Ortalis vetula).
Source: Charles Alexander, Gladys Porter Zoo.
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].
3.6 Numididae – Guineafowl
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.

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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 |