Vermilion flycatcher Temporal range:1.15–0 Ma(present) PreꞒ Ꞓ O S D C P T J K Pg N ↓
Conservation status
Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Passeriformes
Family:	Tyrannidae
Genus:	Pyrocephalus
Species:	P. obscurus
Binomial name
Pyrocephalus obscurus Gould, 1839
Subspecies
See text
Range map[2]   Year-round   Breeding   Nonbreeding

Thevermilion flycatcher (Pyrocephalus obscurus) is a smallpasserine bird in thetyrant flycatcher family found throughout South America and southern North America. It is a striking exception among the generally drab Tyrannidae due to itsvermilion-red coloration. The males have bright redcrowns, chests, and underparts, with brownish wings and tails. Females lack the vivid red coloration and can be hard to identify—they may be confused forSay's phoebe. The vermilion flycatcher's song is apit pit pit pidddrrrreeedrr, which is variable and important in establishing a territory.Riparian habitats and semi-open environments are preferred. As aerialinsectivores, they catch their prey while flying. Their several months-longmolt begins in summer.

Despite being sociallymonogamous, vermilion flycatchers will engage inextra-pair copulation. They also practice within-speciesbrood parasitism, whereby females lay their eggs in the nest of another individual. Females build shallow open cup nests and incubate the brown-speckled whitish eggs. The male feeds the female during incubation. Two broods of two or three eggs are laid in a season lasting from March through June. Once hatched, both males and females feed the chicks, which are ready tofledge after 15 days.

The species was firstdescribed in the late 1830s as a result of the voyages ofCharles Darwin. Thetaxonomy of thegenusPyrocephalus was revised in 2016, which led to the identification of several new species from the vermilion flycatcher'ssubspecies, including the now-extinctSan Cristóbal flycatcher. Populations have declined because ofhabitat loss, though the species remains abundant. The overall population numbers are in the millions, thus theInternational Union for Conservation of Nature considers it a species ofleast concern.

Thetyrant flycatcher family, Tyrannidae, is a group ofpasserine birds present only in theAmericas; its members are generally drab in coloration.^[3] Within it, the subfamily Fluvicolinae comprises thegeneraPyrocephalus,Contopus,Empidonax, andSayornis. They likely share a common ancestor that belonged to the genusContopus orXenotriccus and later diversified. ThePyrocephalus are most closely related toSayornis in terms ofmorphology, but genetic analysis shows they may be more closely related toFluvicola.^[2]

The firstdescription of the vermilion flycatcher was in 1839 byJohn Gould, who created the current genusPyrocephalus, and designated his find asPyrocephalus obscurus. The identification was based on specimens brought back byCharles Darwin on thesecond voyage of HMSBeagle, which lasted from 1831 to 1836.^[4][5] The species was then designated asPyrocephalus rubinus by the English zoologistGeorge Robert Gray in 1840, based on Darwin's specimen taken fromJames Island.^[6][7] A 2016molecular study changed the taxonomy of the species, splitting off several new species and re-designated the original bird asPyrocephalus obscurus.^[8] Before the study it was considered amonotypic genus, but now taxonomists (including theInternational Ornithologists' Union) have elevated three of the vermilion flycatchersubspecies to the rank of species: theDarwin's,San Cristóbal, andscarlet flycatchers.^[9]

The genus namePyrocephalus isGreek and roughly translates to "fire head"^[10] or "flame headed".^[11]: 326 Thespecific epithetobscurus isLatin and means "dark" or "dusky".^[11]: 278 The common name comes from its vibrant coloration and its membership in the flycatcher family, which is reflected in itsinsect-rich diet.^[2]

Before 2016, authors had recognized between 11 and 13 subspecies (sometimes called races). A 2016 molecular study revised that to nine subspecies, made two others their species (P. nanus—Darwin's flycatcher,P. dubius—San Cristóbal flycatcher,P. rubinus—scarlet flycatcher), and determined that another was not valid (P. major). Some works still refer to the vermilion flycatcher asPyrocephalus rubinus, which can lead to confusion with the scarlet flycatcher (also calledPyrocephalus rubinus).^[8][9][12] The vermilion flycatcher likely evolved around 1.15 million years ago (mya), the species on theGalápagos Islands havingsplit off around 0.82 mya. The South American subspecies hadcoalesced by about 0.56 mya, and the North American subspecies had diverged from the South American by 0.25 mya.^[8]

There are nine widely recognized subspecies, which differ primarily in the color andsaturation of the male'splumage and the color and amount of streaking of the female's. The geographic boundaries between some subspecies are not well defined:^[9]

• P. o. obscurus(Gould, 1839)—Thenominate race, which is found in theLima region of western Peru.^[9]
• P. o. mexicanus(Sclater, 1859)—Found from southernTexas in the United States south to central and southern Mexico.^[9] Its upperparts are the blackest of any race, and the male lacks any mottling on the red parts.^[2] This subspecies is named after Mexico.^[11]: 252
• P. o. saturatus(von Berlepsch andHartert, 1902)—Found in northeastern Colombia, western and northern Venezuela, Guyana and northern Brazil.^[9] The female has pink underparts.^[2]Saturatus means "richly colored" in Latin.^[11]: 54
• P. o. blatteus(Bangs in 1911)—Found in southeastern Mexico, Belize and northern Guatemala.^[9] The upperparts are paler, and the underparts more red compared to the nominate race, lacking an orange tinge. It is also smaller than the other Mexican races.^[2]Blatteus means "purple colored" in Latin.^[11]: 73
• P. o. flammeus(van Rossem, 1934)—Found in the southwestern United States and northwestern Mexico.^[9] The upperparts are paler and slightly gray, and the underparts are more orange colored than the nominate race. Males may also have orange mottling in thecrown and breast, whereas the females are less streaky.^[2]Flammeus means "flame colored" in Latin.^[11]: 54
• P. o. ardens(Zimmer, 1941)—Found in northern Peru, in extreme easternPiura,Cajamarca, andAmazonas.^[9] Their coloration has been described as "fiery red". The front of the females crown is slightly pink.^[2]Ardens means "burning" inLatin.^[11]: 54
• P. o. cocachacrae(Zimmer, 1941)—Found from southwestern Peru south to extreme northern Chile.^[9] The male has a brownermantle and less red underparts, while the female has less white underparts, compared to the nominate race. Thetype locality is theCocachacra District in Peru.^[2]
• P. o. piurae(Zimmer, 1941)—Found from western Colombia south to northwestern Peru, it is named for thePiura province in Peru.^{[9][11]: 309}
• P. o. pinicola(Howell, 1965)—Found in eastern Honduras and northeastern Nicaragua.^[9] It is smaller thanP. o. blatteus, and the females have more orange underparts. It preferspinesavanna habitats, which is reflected in the namePinicola: it roughly translates to "pine tree dweller" from Latin.^{[2][11]: 307}

The vermilion flycatcher is a small bird, measuring 13–14 cm (5.1–5.5 in) from tip to tail, around 7.8 cm (3.1 in) from wingtip to body, with a mass between 11 and 14 g (0.39 and 0.49 oz).^[9] Wingspan ranges from 24 to 25 cm (9.4 to 9.8 in).^[13] It is stronglydimorphic. Males are bright red, with contrasting dark brown plumage. Females are drab and have a peach-colored belly with a dark gray upperside. The reddish color varies but can bevermilion,scarlet, or orangish. In males, the crown, chest, and underparts are red. Thelores (region in front of the eyes),nape,ear coverts, wings, upperparts, and tail are all brown to blackish brown. The female has a grayish crown, as well as grayish ear coverts, wings, and tail. Theflight feathers andwing coverts are slightly paler gray, which create a barring effect. Thesupercilium (eyebrow) is whiter. The underparts start white but become light red moving downward. Juveniles of both sexes look similar to adult females; juvenile males have much brighter red underparts, whereas juvenile females have yellowish underparts. Plumage appears constant throughout the year for both adult sexes and for juveniles.^[2] They have a slightcrest, which can be raised when needed.^[14] Males are not easily mistaken for other species, but the drab females may be confused with theSay's phoebe.^[2]

Worn feathers are replaced bymolting, which takes between 62 and 79 days and begins in July, lasting until September. Many vermilion flycatchers molt only after completing theirmigration to warmer regions. The molt is fairly slow compared to that of other families, as quick molting creates poor feathers and interrupts flight, which is untenable for an aerially feeding species. A 2013 study determined thatmonsoon rain patterns do not affect molting, as had been previously expected. Instead,latitude-based temporal effects are more important in timing the molt.^[15]

OrnithologistDavid Sibley describes the perchingsong as apit pit pit pidddrrrreedrr,^[16] whereas theCornell Lab of Ornithology describes it as aching-tink-a-le-tink, with an emphasis on the last syllable.^[2] The flight song is given by males who fly high above the canopy, and is described as apt-pt-pre-ee-een by the Cornell Lab. The regular song may also be given as a slower chatter call. Other noises include apees noise that is given as a call. Apeent is given while foraging, but a more aggressive variety is also used between males. As a lead-up tocopulation, the female may give atjee-tjee-tjee call.^[2][16]

The call of the vermilion flycatcher is important in establishing its territory. Males make only a single song while perching, but can alter the song to convey different intentions. The male's song consists of a variable first part, and a second part with four elements. The first part of the song gets longer after nest construction, and before dawn. The first part encodes information regarding the quality of the male to potential mates and the level of threat to other males—which is related to the intensity with which males defend their territory.^[17] Females do not usually sing.^[18] Urbannoise pollution appears to affect songs; a population inMexico City was found to sing louder and longer songs as ambient noise increased.^[19] Singing is not heard year-round; populations in Arizona and Texas sing only from late February through July.^[2]

Non-vocal sounds include snapping the beak, by males between songs and by females while watching male courtship flights. Their wings may also be used to create a whirring noise while perch hopping or during territorial displays, though this is practiced infrequently.^[2]

Their range includes almost all of Mexico, extending north into thesouthwestern United States, and south to scattered portions ofCentral America, and parts of northwestern and centralSouth America. It has ranged as far north as Canada.^[20] North American populations are generallyresident, migrating only at the edge of the range. South American populations, especially those further south, may make long migrations to the northernmost parts of theBrazilian Amazon. This reflects a tendency to overwinter in areas where the temperature does not go below −1 °C (30 °F). Migrations may extend up to 4,000 km (2,500 mi). North American populations generally migrate by late August and return between February and April.^[2] Their ability to migrate likely aided their wide colonization of the Americas.^[8]

Vermilion flycatchers prefer somewhat open areas and are found in trees or shrubs in savanna, scrub, agricultural areas,riparian woodlands, and desert as well, but usually near water.^[20] They range up to elevations of 3,000 m (9,800 ft). A study in Arizona found that their preferred breeding range includedcottonwood ormesquite tree canopies, althoughFremont cottonwoods were not favored.Goodding's willow was preferred as a nesting site where found. Understory plants mainly consisted of invasiveCynodon dactylon grass.^[2]

Vermilion flycatchers are generally solitary, though they may form small flocks of not more than five individuals during winter. They spend most of their time in trees perching, landing on the ground only rarely to catch insects. They do not generally hop, preferring to fly to get around, andglide only infrequently.^[2]

Vermilion flycatchers are sociallymonogamous, but engage inextra-pair copulation. Both males and females will breed with individuals other than their monogamous mate. A 2002 study found that 11% of offspring were from extra-paternal copulation. They also practice intra-specificbrood parasitism, where females lay their eggs in the nest of another vermilion flycatcher. Between 9.5 and 19% of offspring were the result of brood parasitism. Females often spend long times away from the nest, which enables others to lay eggs in their nests while absent. This may simply be a form of parasitism, wherein the pair whose nest is being parasitized gains no benefit. But for promiscuous birds, this may provide somegenetic fitness. For a male mating with many females, and those females laying many of their eggs outside of their nest, this increases the odds that a promiscuous male's offspring are laid in his nest. This allows the male to outsource the energy-intensive process of egg-laying away from his mate and allows a female to outsource the process of raising her chick. This is supported by examples of males letting females parasitize their nests in exchange for copulation.^[18] The flycatcher is a frequent victim of brood parasitism bybrown-headed cowbirds.^[2][21] Males establish and aggressively defend territory during breeding season. While defending, males raise their crest and fan their tail out. They will also pump their tail and snap their beak. Males will chase other males out of their territory by flying after them. Once nestlings are present, they will also chase off other bird species. Courtship involves the male fluffing his crest and chest feathers, fanning the tail, and engaging in a fluttery flight while singing to a female.^[2][14]

Males then choose a nesting site, and females begin construction. The vermilion flycatcher'snest is a shallow cup made of small twigs and soft materials, lined with feathers; the nest's rim is often covered withlichen. Spiderwebs may be used to bind the nest together. Females use a rocking motion of their body to shape the nest during construction. Even after construction, the female will continue to add insulating material, such as plant materials, hair, fur, or man-made materials like string. Average nest sizes are about 64–76 mm (2.5–3.0 in) across, 25–51 mm (0.98–2.01 in) tall, and with a cup depth of less than 25 mm (0.98 in).^[2] Nests are typically located within 6 ft (1.8 m) of the ground; the nest is placed in the horizontal fork of a tree branch. About 12% of nests are reused, and old nests may be raided for materials for new nests.^[22]

Egg-laying begins in March and runs through June; eggs are laid once a day in the early morning. Eggs are ovate and approximately 17 mm (0.67 in) × 13 mm (0.51 in). They average 1.6 g (0.06 oz) in weight, which is about 11% of the female's body mass. They are a dull whitish color and have large brown splotches in a wreath pattern on the larger end, though egg coloration varies. Spotting may be more intense, and the base color may be creamy or even tan or brown. Clutches usually contain two or three eggs but may occasionally include up to four.^[2] Eggs are incubated solely by the female for 13–15 days. The male feeds the female while she incubates eggs, though females never beg for food. Feedings are sometimes followed by copulation. The female is very attentive while in the nest. The young arealtricial, meaning they are incapable of fending for themselves after birth. Both parents feed the chicks, although the male may tend to fledglings while the female builds another nest.^[2][23] Nests may be reused during the same season, but this is uncommon; one study found that only 12% of nests were reused, and only if they had been successful in raising a brood. Re-using nests saves time and energy but perhaps at the cost of a higher parasite load.^[24] Nestlings open their eyes four days after hatching. The young are ready toleave the nest 15 days after hatching. All young generally fledge on the same day, although some may leave a day earlier. If the nest is disturbed, nestlings older than 11 days will abandon the nest prematurely. There are usually two broods per year, although three are possible.^[2][23]

• 
  Male at nest
• 
  Nestlings, who have yet to hit their first juvenile molt
• 
  Juvenile maleP. o. obscurus inLima,Peru, showing off the lighter juvenile plumage

The vermilion flycatcher feeds mostly on insects such asflies,grasshoppers andbeetles—though the exact composition of the diet is poorly studied.^[25] These are usually taken in mid-air, after a short sally flight from a perch.^[2][26] It is anopportunistic feeder, and has been observed eating smallfish, though it is not known to eat plant material.^[2][27] Bees may also be taken as forage. Non-digestible insect parts are regurgitated as pellets.^[23] While waiting for insects, they will sit on thin branches and pump their tail up and down.^[14] While active, about 90% of their day is spent perching, and only 4–11% is spent chasing prey. Once they have spotted prey, they jump up from their perch and give chase. If insects are missed on the first attempt, the bird is capable of quite a nimble flight to catch them. Once caught, the insects may be beaten before being swallowed whole. Occasionally, insects will be caught on the ground. Otherwise, most prey is caught within 3 m (9.8 ft) of the ground and rarely above water.^[2]

The predators of the vermilion flycatcher are not well known. Unusual reports of predation include by ascrub-jay, and a group of live nestlings eaten byfire ants. The oldest recorded individual lived to five and a half years, but otherwise, lifespan data is lacking, as is data about mortality causes. Yearly nesting success (the percentage of laid eggs which were raised to fledglings) in a Texas study varied from 59 to 80%. Half of the lost nests contained eggs and half contained young. The causes of failure included nest abandonment and egg infertility. A similar study in Ecuador showed success rates from 20 to 59%.^[2]

Severalectoparasites affect the species,Dermanyssus mites being common. A 2008 study found that mites did not greatly affect nesting success, nor did nesting reuse greatly enhance mite populations. Fledglings were successfully raised even in infested nests, while nearby, un-infested nests, did not raise any fledglings. Though not directly parasitic,flies may lay their eggs in nests, providing a home for theirlarva.^[24]

Although the diseases of the vermilion flycatcher are not well studied, the diseases of the birds of the Galapagos are known, and these diseases may have affected the closely related San Cristóbal flycatcher. Introduced and destructive diseases includeavian malaria,Marek's disease,Newcastle disease, and many others.^[28]Avian pox viruses and crop canker (caused byTrichomanes gallinae) may have directly contributed to the extinction of the Galapagos sister species.^[29]

The vermilion flycatcher is a favorite withbirders, but it is not generally kept inaviculture as the males tend to lose their vermilion coloration when in captivity.^[30] This is likely a diet-based effect, as maintaining bright red coloration in birds requires substantial quantities of yellow precursorzeaxanthin molecules which are thenmetabolized into red pigment. This acts as an indicator of genetic fitness to potential mates, as a bright male is using his diet-based coloration to show off his ability to survive and catch food.^[31][32]

TheTucson Bird Alliance (formerly Tucson Audubon Society) ofTucson, Arizona, publishes an eponymous journal named for the Vermilion Flycatcher.^[33]

Because of its enormous range and sizable population—with a population estimated ranging between 5,000,000 and 50,000,000 individuals—the vermilion flycatcher is listed as a species ofleast concern by theInternational Union for Conservation of Nature.^[1] Its overall numbers are declining; between 1966 and 2007, populations declined an average of 1.7% per year. Texas populations are declining faster, at 2.6% per year. Once prolific in southernCalifornia, it has become increasingly rare, although it is expanding into new areas such asFlorida andOklahoma. Arizona populations are increasing, at a rate of 2.2% per year. Vermilion flycatchers have adapted to human structures by increasingly nesting in parks and golf courses. However, the increased productivity of these areas may be outweighed by an increased presence of brown-headed cowbirds—whose parasitic young leave less food for flycatcher young—as well as increased nest predation.Habitat destruction is a major concern for the flycatcher, especially inriparian areas. A prime example is along theLower Colorado River Valley, where changes in water management combined with the destruction of cottonwood-willow riparian habitat have led to the loss of almost all breeding and foraging areas.^[2]

The San Cristóbal flycatcher, which was once considered part of the species, was endemic to the Galápagos Islands but went extinct sometime between 1987 and 2012.^[34][35] The Darwin's flycatcher, which was also once part of the species, is considered anear-threatened species by theInternational Union for Conservation of Nature.^[36]

Wikimedia Commons has media related toPyrocephalus obscurus.

Wikispecies has information related toPyrocephalus obscurus.

• "Vermilion flycatcher media".Internet Bird Collection.
• Species account—Cornell Lab of Ornithology
• Vermilion flycatcher—Pyrocephalus rubinus—USGS Patuxent Bird Identification InfoCenter
• Stamps at bird-stamps.org
• Vermilion flycatcher photo gallery at VIREO (Drexel University)
• Interactive range map ofPyrocephalus rubinus atIUCN Red List

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• Taxa named by John Gould

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