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Pharomachrusquetzals

By Nathan Hollars

Diversity

Genus Pharomachrus consists of five species. Pharomachrus mocinno, commonly known as the resplendent quetzal, is the most well-known, named by naturalist Pable de la Llave. Within this species, there are two subspecies: P. m. mocinno and P. m. costaricensis. The second species in Pharomachrus, P. antisianus, is commonly known as the crested quetzal. The third species, P. auriceps, is commonly known as the golden-headed quetzal due to its distinctly golden head. This species consists of the subspecies P. a. auriceps and P. a. hargitti. The fourth species, P. pavoninus, is commonly known as the pavonine quetzal or peacock trogon, distinguishable by its red bill. Finally, P. fulgidus, commonly known as the white-tipped quetzal, has two subspecies: P. f. fulgidus and P. f. festatus. All Pharomachrus quetzals are distinct due to their combination of vibrant, iridescent colors and their long wing/tail coverts. (Rafael and Sittler, 2019; Solórzano and Oyama, 2001)

Geographic Range

Respledent quetzals (P. mocinno) and their subspecies are found throughout southern Mexico, ranging from Guatemala to Panama. They are mainly concentrated in Guatemala and Honduras, with additional populations in Nicaragua, Costa Rica, and Panama. Crested quetzals (P. antisianus) along with golden-headed quetzals (P. auriceps) and their subspecies are found throughout the upper regions of South America, ranging from Venezuela, through Colombia, Ecuador, and Peru to Bolivia. Pavonine quetzals (P. pavoninus) are found throughout northern South America, ranging from Colombia through Venezuela, Ecuador, Peru, and Bolivia to Brazil. White-tipped quetzals (P. fulgidus) and their subspecies are found in a small region of northern Colombia and small regions of northern Venezuela. All species and subspecies of Pharomachrus are native residents to their respective regions - except white-tipped quetzals, which were introduced to Venezuela by humans. (BirdLife International, 2016; Lohnes and Greeney, 2008; Rafael and Sittler, 2019)

Habitat

Members of Pharomachrus typically live in mountainous/forested areas, specifically in the canopies and sub-canopies of humid montane forests and cloud forests. They prefer regions with ravines as well as cliffs covered with vegetation. Various types of large oak trees (ranging from 100-150+ ft tall) form the forests' canopies. Smaller alders and various laurels, both of which are an important source of fruit for resident birds, also comprise the forests.

Depending on the species, Pharomachrus can be found at elevations ranging between 900m - 3,200m. While most members of the genus inhabit the highlands of their respective regions, P. pavonius occupies the lowlands. Pharomachrus members nest/shelter in cavities within dead tree trunks, and P. moccino have been observed modifying sites abandoned by woodpeckers. Members Pharomachrus will often seek out clearings within and beyond the forests to nest and forage. (BirdLife International, 2016; Lohnes and Greeney, 2008; Rafael and Sittler, 2019; Skutch, 1944)

Systematic and Taxonomic History

The relationships between the Pharomachrus genus and other genera in order Trogoniformes are well-understood with multiple, ongoing investigations attempting to better understand their relations. Genus Pharomachrus is most closely related to genus g. Euptilotis. Within order Trogoniformes, both genera are considered to be sisters to the rest of the trogons.

Genus Pharomachrus has no previously-used synonyms; however, each individual species has at least one synonym which was previously used in published literature. Currently, there is not much understanding of the relationships between each species and respective subspecies within Pharomachrus, as there have only been a few studies conducted. Some researchers suggest that subspecies P. m. mocinno and P. m. costaricensis should be considered their own species instead of subspecies. (Johansson and Ericson, 2004; Lepage, 2014; Moyle, 2001; Solórzano and Oyama, 2001)

  • Synonyms
    • Trogon antisiensis
    • Pharomachrus auriceps xanthogaster
    • Pharomachrus xanthogaster
    • Pharomachrus auriceps heliactin
    • Pharomachrus pavoninus heliactin
    • Pharomachrus festatus
    • Pharomachrus pavoninus hargitti
    • Pharomachrus xanthogaster hargitti
    • Pharomacrus costaricensis
    • Pharomachrus pavoninus viridiceps

Physical Description

All species in genus Pharomachrus are known for their iridescent/golden-green feathers covering their back, throat, and upper wing coverts. They also have characteristically red feathers covering their bellies as well as their breasts. They have black primaries and secondaries on their wings, along with short yellow/yellow-orange bills and brown/dull green feet.

Male and female members of P. mocinno are sexually dimorphic, with males having distinctly long, iridescent, green tail feathers along with a red breast and a crest on their head. Females have a short white and black tail with a chevron pattern and a gray breast. Juveniles are brown with mottled scapulars and coverts. The upper portion of their breasts are brown and the lower portion is white. Their bellies are white with greyish brown coloration and they have black bills and lead-colored feet.

P. auriceps has a distinct golden brown/bronze head. Females have a brown head and a brownish breast and bill.

P. fulgidus is distinct in having a black upper tail with a white tip. Females differ in having a brown head, breast, and beak.

P. antisianus males have a tuft of feathers forming a crest protruding out of their forehead, similar to P. mocinno, but distinguishable by the lack of long tail feathers and red eyes. P. antisianus females have a brown head and belly with a mostly black tail.

P. pavoninus males are distinguishable by a red bill whereas females have a brown/gray head and bill. (Cara Grace, 2016; Rafael and Sittler, 2019)

  • Sexual Dimorphism
  • sexes colored or patterned differently
  • male more colorful

Reproduction

Pharomachrus species typically breed at high elevations where the males find and maintain monogamous relationships. P. mocinno males will use a coouee whistle as a mating call, in which notes rise like a police siren. Before calling out to mates, males will establish a territory and use a two-note whistle, during which they hold their head high and raise their chests with their bill slightly open and their breast puffed out. They repeat this call every 8-10 minutes and use it to advertise their territory. Once a breeding pair is formed, they remain together during the next three breeding seasons, and most likely for the rest of their life. Other Pharomachrus species mating systems are understudied. (LaBastille, et al., 1972; Rafael and Sittler, 2019)

Pharomachrus species are seasonal breeders with breeding seasons beginning in March or May (depending on location) and lasting 1-2 months. After finding a mate, the male and female build nests together, taking turns to widen cavities in dead tree trunks. Females typically lay a clutch size of two sky-blue eggs, with some clutches being laid in different nests and some nests being used for multiple clutches. The subsequent clutch is laid after the previous one is fledged. All Pharomachrus species seem to follow a very similar nest-building, clutch-laying process. A notable exception is that the two eggs in the clutch of P. pavoninus females are sky-blue in color with a few small brown speckles. (Daniel, 2007; Lohnes and Greeney, 2008; Paulo and Oscar, 2015; Rafael and Sittler, 2019; Wheelwright, 1983)

In Pharomachrus, both the male and female participate in building and carving out a site for the nest. Once the clutch is laid, both males and females share incubation shifts in which females incubate at night and males incubate in the early afternoon. Chicks hatch 18-19 days after being laid. After their eggs hatch, parents take turns remaining in the nests and feeding the young. Nests are attended to for 60-95% of the day during the offsprings' first 6-8 days and then 20-40% later in the nesting period. This decline in brooding is presumed to be associated with the chicks having acquired the ability to thermoregulate. Towards the end of the nesting period, parents perch nearby in order to attend to the nest. The amount of time spent by both males and females brooding, attending the nest, and bringing food varies between nests/pairs. (Daniel, 2007; Lohnes and Greeney, 2008; Paulo and Oscar, 2015; Rafael and Sittler, 2019; Wheelwright, 1983)

Lifespan/Longevity

Due to the elusive nature of Pharomachrus, little is known about their lifespans. A major limiting factor on their lifespans, however, is habitat degradation, largely due to anthropomorphic activities causing large portions of cloud forests to be deforested. The unavailability of nesting sites is a consequence of habitat destruction, which ultimately impacts Pharomachrus population sizes. In the wild, predators include gray squirrels, ornate hawk-eagles, margays, and green toucanets.

Pharomachrus species have also been poached and trapped for many years. Their feathers were extremely valuable to the Mayans and Aztecs, and they continue to be valuable today. Live specimens are often caught and sold. (Cara Grace, 2016; Rafael and Sittler, 2019)

Behavior

Pharomachrus are largely solitary birds until the breeding season, during which they will find and stay with a mate. They will remain with that mate for the next few breeding seasons, at least. P. mocinno has been described as having an apprehensive disposition and calm manner, perching motionlessly on a branch for long periods of time. This perching behavior is assumed to be exhibited throughout the genus. During the mating season, however, P. mocinno is much more active, using a variety of vocalizations. It should be noted that males have distinct, unique calls. Additionally, aerial displays are exhibited, during which the male's tail feathers are fully exhibited. Breeding season is the only time where P. mocinno individuals migrate short distances to lower elevations in search of nest sites and food availability. Females have been seen attacking other quetzals trespassing on their territory. After breeding season, males will molt and their tail feathers will begin to grow again in time for the next mating season. (Cara Grace, 2016; LaBastille, et al., 1972; Rafael and Sittler, 2019)

Communication and Perception

P. mocinno uses many different vocalizations. A two-note whistle is used by males to advertise territory to other males. A recognition call described as a wac-wac sound is also used between mates. When alarmed, P. mocinno uses a monotone note call along with a tail flick and opening their tail feathers like a fan. As a whole, however, species in genus Pharomachrus are understudied when it comes to communication and perception. (Daniel, 2007; LaBastille, et al., 1972)

Food Habits

Pharomachrus adults are primarily frugivorous. However, chicks are described as having an omnivorous diet consisting of insects and small vertebrates, such as lizards and fruits. P. mocinno adults have been observed feeding on fruits from 15 different plant species, with 40% of them being laurels. Other consumed plant species included members of families Theaceae (tea family), Myrsinaceae (myrsine family), Araliaceae (ginseng family), Verbenaceae (verbena family), Solanaceae (nightshades), Myrtaceae (myrtle family), Melasomataceae (melastomes), and Mora. These plant families consist of shrubs, trees, herbs, and general flowering plants. It is estimated that Pharomachrus species annually feed on a total of ~41 species. They primarily consume large drupes along with smaller, seeded berries. Chicks' diets change after their first 10 days, with 24% of their initial diet consisting of fruits and 76% being insects/small vertebrates, and after the tenth day, their diet consists of 72% fruits and 28% insects/small vertebrates. Members of genus Pharomachrus are important seed dispersers, as they have a specialized behavior in which they regurgitate seeds far from the original tree source. The diet of P. auriceps chicks is seemingly less varied in terms of animal matter consumed, as it appears that they are primarily insectivores. P. pavoninus chicks are unique in that they have been observed consuming frogs. (Avila H., et al., 1996; Cara Grace, 2016; Daniel, 2007; Lohnes and Greeney, 2008; Rafael and Sittler, 2019; Wheelwright, 1983)

Predation

Pharomachrus species experience little predation, although there have been a few recorded cases. This is in part due to their size, but also their green coloration. The green plumage provides camouflage and is particularly effective in the rain because their feathers show very little iridescence, allowing them to blend in with wet vegetation. Pharomachrus species' motionless perching behavior also plays a role in protecting them from predators because they will sit with their red chest facing away from potential danger. This, in conjunction with their green coloration, makes them difficult to spot. Predation primarily takes place in the form of stealing eggs from the nest and attacks on Pharomachrus juveniles. (Cara Grace, 2016; LaBastille, et al., 1972; Rafael and Sittler, 2019; Wheelwright, 1983)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Pharomachrus species are predators to a number of different insects and some small vertebrates such as lizards. Additionally, P. pavoninus is a predator to frogs. The eggs and juveniles of Pharomachrus species are a food source to a number of different species. The primary ecological role of Pharomachrus species, however, is seed dispersal. They typically regurgitate seeds about 15-30 minutes after ingestion and only spend short times at fruiting trees. These factors lend a very high chance that seeds will be regurgitated far away from the original tree, leading to effective dispersal. Pharomachrus species also play a key role in the amount of available resources for other species. They are highly selective about the fruits they consume, only selecting fruits that are large and ripe in order to gain the most nutrients for the associated energy cost. This leaves a large selection of small fruits and a small selection of high-quality fruits available for other species. This combination of selectiveness and diet specialization makes Pharomachrus species important for the ecological health of cloud forests. (Avila H., et al., 1996; Rafael and Sittler, 2019; Wheelwright, 1983)

  • Ecosystem Impact
  • disperses seeds

Economic Importance for Humans: Positive

The feathers of Pharomachrus species were extremely valuable to the Mayan culture because they were used to adorn clothing. The feathers are still highly valuable today; they are sold to tourists and museum collectors along with dried specimens of the birds themselves and some sales of living individuals.

In Guatemala, quetzals are the national emblem, holding great importance to the country's culture. P. mocinno is considered a flagship species, meaning they attract ecotourists wanting to observe the species in protected natural areas. This is considered an important source of income.

There have been many studies on Pharomachrus species, with a number of research projects done on P. mocinno. These studies have provided educational benefits about the birds as well as their cloud forest ecosystems. (Cara Grace, 2016; Rafael and Sittler, 2019; Skutch, 1944)

  • Positive Impacts
  • body parts are source of valuable material
  • ecotourism
  • research and education

Economic Importance for Humans: Negative

There are no known adverse effects of Pharomachrus species on humans.

Conservation Status

Pharomachrus species are largely considered to be of Least Concern when it comes to conservation; however, P. mocinno is considered Near Threatened. This status is mostly due to habitat destruction as well as trapping/poaching. There are a number of ongoing conservation efforts, especially when concerning P. mocinno. Some conservation areas have been set up with protected populations of P. mocinno in order to help preserve them and the cloud forests that they inhabit. There is additional research, monitoring, and planning also occurring in order to better conserve this species since P. mocinno fits the description of being potentially Vulnerable due to their restricted diets and low population densities. (BirdLife International, 2016; Rafael and Sittler, 2019; Wheelwright, 1983)

  • IUCN Red List [Link]
    Not Evaluated

Other Comments

The name "Pharomachrus" comes from the ancient Greek words pharos and makros, meaning "mantle" and "long" respectively. This name was likely given in reference to the birds' distinctly long tails and wing coverts. Pharomachrus species are culturally important, dating back to the Mayans and Aztecs, and are associated with Quetzalcoatl, a diety. Overall, these birds' feathers were considered to be symbolic/sacred and are still sought after today. (Rafael and Sittler, 2019)

Contributors

Nathan Hollars (author), Colorado State University, Sydney Collins (editor), Colorado State University.

Glossary

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

choruses

to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

insectivore

An animal that eats mainly insects or spiders.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males

migratory

makes seasonal movements between breeding and wintering grounds

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

References

Avila H., M., V. Hernandez O., E. Velarde. 1996. The Diet of Resplendent Quetzal (Pharomachrus Moncinno mocinno: Trogonidae) in a Mexican Cloud Forest. Biotropica, Vol 28. No. 4: pg. 720-727. Accessed February 10, 2022 at https://www.jstor.org/stable/2389058.

BirdLife International, 2016. "Pharomachrus mocinno" (On-line). The IUCN Red List of Threatened Species. Accessed February 10, 2022 at https://www.iucnredlist.org/species/22682727/92958465.

Cara Grace, T. 2016. Birds of a Feather: Exploring the Acquisition of Resplendent Quetzal (Pharomachrus mocinno) Tail Coverts in Pre-Columbian Mesoamerica. Human Ecology, Vol. 44: 399-408. Accessed February 17, 2022 at https://link.springer.com/article/10.1007/s10745-016-9827-8#citeas.

Daniel, J. 2007. Nesting Behavior and Nestling Care of the Pavonine Quetzal (Pharomachrus pavoninus). Wilson Journal of Ornithology, Vol.119 (3): 458-463. Accessed February 19, 2022 at https://bioone.org/journals/the-wilson-journal-of-ornithology/volume-119/issue-3/06-138.1/Nesting-Behavior-and-Nestling-Care-of-the-Pavonine-Quetzal-span/10.1676/06-138.1.short.

Johansson, U., P. Ericson. 2004. A re-evaluation of basal phylogenetic relationships within trogons (Aves: Trogonidae) based on nuclear DNA sequences. Journal of Zoological Systematics and Evolutionary Research, Volume 43, Issue 2: Pages 166-173. Accessed February 06, 2022 at https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0469.2004.00292.x.

LaBastille, A., . Allen, L. Durrell. 1972. Behavior and Feather Structure of the Quetzal. The Auk, Volume 89, Issue 2: Pages 339–348. Accessed February 06, 2022 at https://academic.oup.com/auk/article/89/2/339/5209322.

Lepage, D. 2014. "Avibase - The World Bird Database" (On-line). Accessed March 27, 2022 at https://avibase.bsc-eoc.org/avibase.jsp.

Lohnes, R., H. Greeney. 2008. Brooding behaviour and nestling description of Golden- headed Quetzal Pharomachrus auriceps. Cotinga, Volume 30: Pages 47-50. Accessed February 06, 2022 at https://www.semanticscholar.org/paper/Brooding-behaviour-and-nestling-description-of-Lohnes-Greeney/eca662f1d2425c87c689408a0ffc53692c5ca63d.

Moyle, R. 2001. Phylogeny and biogeographical history of Trogoniformes, a pantropical bird order. Biological Journal of the Linnean Society, Volume 84, Issue 4: Pages 725–738. Accessed February 06, 2022 at https://academic.oup.com/biolinnean/article/84/4/725/2701424.

Paulo, ., A. Oscar. 2015. The Nest and Eggs of the White-tipped Quetzal (Pharomachrus fulgidus) from the Sierra Nevada de Santa Marta, Northern Colombia. The Wilson Journal of Ornithology, Vol.127 (1): 145-148. Accessed February 19, 2022 at https://bioone.org/journals/the-wilson-journal-of-ornithology/volume-127/issue-1/14-018.1/The-Nest-and-Eggs-of-the-White-tipped-Quetzal-Pharomachrus/10.1676/14-018.1.short.

Rafael, P., B. Sittler. 2019. "Hal theses" (On-line). Acoustic behavior and ecology of the Resplendent Quetzal Pharomachrus mocinno, a flagship tropical bird species. Accessed February 06, 2022 at https://tel.archives-ouvertes.fr/tel-02048769.

Skutch, A. 1944. Life History of the Quetzal. The Condor, Vol. 46: pg. 213-235.

Solórzano, S., A. Baker, K. Oyama. 2001. Conservation Priorities for Resplendent Quetzals Based on Analysis of Mitochondrial DNA Control-Region Sequences. The Condor, Vol. 106 No. 3: pg. 449-456. Accessed February 06, 2022 at https://www.jstor.org/stable/4151043.

Solórzano, S., G. Mara, K. Oyama. 2009. Genetic diversity and conservation of the Resplendent Quetzal Pharomachrus mocinno in Mesoamerica.. Revista mexicana de biodiversidad, Volume 80 (1): Pages 241-248. Accessed February 06, 2022 at https://www.semanticscholar.org/paper/Genetic-diversity-and-conservation-of-the-Quetzal-Sol%C3%B3rzano-Garc%C3%ADa-Ju%C3%A1rez/c726620452e2f8c4ac79fc2bda062f718a7cc2a6.

Solórzano, S., K. Oyama. 2001. Morphometric and molecular differentiation between quetzal subspecies of Pharomachrus mocinno (Trogoniformes: Trogonidae). Revista De Biología Tropical, Vol.58 (1): pg. 357-371. Accessed February 06, 2022 at https://revistas.ucr.ac.cr/index.php/rbt/article/view/5215.

Wheelwright, N. 1983. Fruits and the Ecology of Resplendent Quetzals. The Auk, Vol. 100: pg. 286-301. Accessed February 06, 2022 at https://www.jstor.org/stable/4086524?origin=pubexport.

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