Galliformes: Species, Varieties, and Behavior


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Galliformes: 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].

Schematic illustration of galliformes cladogram displaying the relationships among families of the Galliformes. The taxonomic and phylogenetic relationships within this order remain debated and ever evolving.

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
Photo depicts maleo (Macrocephalon maleo).

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
Photo depicts cracidae. (a) Wattled curassow (Crax globulosa) hen.

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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Nov 10, 2022 | Posted by in GENERAL | Comments Off on Galliformes: Species, Varieties, and Behavior

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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