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The red warbler (''Cardellina rubra'') is a small
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
bird of the New World warbler family Parulidae
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the highlands of Mexico, north of the Isthmus of Tehuantepec. It is closely related to, and forms a superspecies with, the
pink-headed warbler The pink-headed warbler (''Cardellina versicolor'') is a small passerine bird found in the southwestern highlands of Guatemala and the central and southeastern highlands of the Mexican state of Chiapas. The adult is primarily red, with a silvery ...
of southern Mexico and Guatemala. There are three
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
, found in disjunct populations, which differ in the color of their ear patch and in the brightness and tone of their body plumage. The adult is bright red, with a white or gray ear patch, depending on the subspecies; young birds are pinkish-brown, with a whitish ear patch and two pale . Breeding typically occurs between February and May. The female lays three or four eggs in a domed nest, which she builds on the ground. Though she alone incubates the eggs, both sexes feed the young and remove
fecal sac A fecal sac (also spelled faecal sac) is a mucous membrane, generally white or clear with a dark end, that surrounds the feces of some species of nestling birds. It allows parent birds to more easily remove fecal material from the nest. The nes ...
s from the nest. The young
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
within 10–11 days of hatching. The red warbler is an
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
, gleaning primarily in understory shrubs. Although this bird is considered to be a least-concern species according to the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN), its numbers are thought to be declining due to
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
.


Taxonomy

English jeweler and naturalist William Bullock and his son traveled to Mexico soon after its
independence Independence is a condition of a person, nation, country, or state in which residents and population, or some portion thereof, exercise self-government, and usually sovereignty, over its territory. The opposite of independence is the statu ...
, spending six months in 1823 collecting archaeological artifacts and many bird and fish species new to science. The bird specimens were given to naturalist
William John Swainson William John Swainson FLS, FRS (8 October 1789 – 6 December 1855), was an English ornithologist, malacologist, conchologist, entomologist and artist. Life Swainson was born in Dover Place, St Mary Newington, London, the eldest son of ...
, their countryman, to formally
describe Shneur Hasofer is a Hasidic musician known as DeScribe. Hasofer's musical style has been characterized as "Hasidic hip-hop," "Hasidic rap" and "Hasidic R&B". Background Hasofer was born to a Chabad Hasidic family in Melbourne, Australia. Hasof ...
, which he did in 1827. Among these was the red warbler, which was assigned to the genus '' Setophaga'', as ''Setophaga rubra''. Over the next half century, other authorities moved it to ''
Cardellina ''Cardellina'' is a genus of passerine birds in the New World warbler family Parulidae. The genus name ''Cardellina'' is a diminutive of the Italian dialect word ''Cardella'' for the European goldfinch. The genus was introduced by the French na ...
'', with the
red-faced warbler The red-faced warbler (''Cardellina rubrifrons'') is a species of New World warbler. Mature red-faced warblers are small birds, long. They are light gray on top with a white rump and a white underside. The face, neck, and upper breast are all b ...
, and to the widespread tropical warbler genus ''
Basileuterus ''Basileuterus'' is a genus of New World warblers, best represented in Central and South America. This is one of only two warbler genera that are well represented in the latter continent. Some species formerly considered in this genus are now pla ...
'', as well as to the
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
warbler genus '' Sylvia'' and the Old World tit genus '' Parus''. In 1873, English naturalists
Philip Lutley Sclater Philip Lutley Sclater (4 November 1829 – 27 June 1913) was an English lawyer and zoologist. In zoology, he was an expert ornithologist, and identified the main zoogeographic regions of the world. He was Secretary of the Zoological Society o ...
and Osbert Salvin moved the species to the genus '' Ergaticus'', where it remained for more than a century. The red warbler forms a superspecies with the pink-headed warbler of
Chiapas Chiapas (; Tzotzil language, Tzotzil and Tzeltal language, Tzeltal: ''Chyapas'' ), officially the Free and Sovereign State of Chiapas ( es, Estado Libre y Soberano de Chiapas), is one of the states that make up the Political divisions of Mexico, ...
and Guatemala. Despite their disjunct ranges and considerably different plumages, the two have sometimes been considered conspecific. Conversely, it has also been suggested that the red warbler should be split into a northern gray-eared species (''C. melanauris'') and a southern white-eared species (''C. rubra''). A comprehensive 2010 paper by Irby Lovette and colleagues analyzing
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
and nuclear DNA of the New World warblers found that the red and
pink-headed warbler The pink-headed warbler (''Cardellina versicolor'') is a small passerine bird found in the southwestern highlands of Guatemala and the central and southeastern highlands of the Mexican state of Chiapas. The adult is primarily red, with a silvery ...
s are each other's closest relative and that their common ancestor diverged from a lineage that gave rise to the red-faced warbler. The authors recommended moving the red and pink-headed warblers to the genus ''Cardellina'', a suggestion which has been adopted by the International Ornithologists' Union (IOC). There are three subspecies, which differ slightly in appearance: * ''C. r. rubra'', described by Swainson in 1827, has white ear patches and is found from southern
Jalisco Jalisco (, , ; Nahuatl: Xalixco), officially the Free and Sovereign State of Jalisco ( es, Estado Libre y Soberano de Jalisco ; Nahuatl: Tlahtohcayotl Xalixco), is one of the 31 states which, along with Mexico City, comprise the 32 Federal En ...
and southern Hidalgo to
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
. * ''C. r. melanauris'', originally described and named by American ornithologist
Robert Thomas Moore Robert Thomas ("R. T.") Moore (June 24, 1882 – October 30, 1958) was an American businessman, ornithologist, philanthropist, the founder and Editing, editor-in-chief of the Borestone Mountain Poetry Awards. In his obituary, Lionel Stevenson wrote ...
in 1937 as ''Ergaticus ruber melanauris'', has dark gray ear patches and somewhat more scarlet upperparts than ''C. r. rubra''. The subspecific name is derived from the Ancient Greek ''melan-'' "black/dark", and Latin ''auris'' "ear". It is found from southwestern
Chihuahua Chihuahua may refer to: Places *Chihuahua (state), a Mexican state **Chihuahua (dog), a breed of dog named after the state **Chihuahua cheese, a type of cheese originating in the state **Chihuahua City, the capital city of the state **Chihuahua Mun ...
to northern Nayarit. * ''C. r. rowleyi'' was originally described and named by R. T. Orr and J. D. Webster in 1968 as ''Ergaticus ruber rowleyi'', in honor of J. Stewart Rowley, a research associate at the
California Academy of Sciences The California Academy of Sciences is a research institute and natural history museum in San Francisco, California, that is among the largest museums of natural history in the world, housing over 46 million specimens. The Academy began in 1853 ...
. It has white ear patches and ruby-red upperparts (brightest of the three subspecies), and is found in the
Sierra Madre del Sur The Sierra Madre del Sur is a mountain range in southern Mexico, extending from southern Michoacán east through Guerrero, to the Isthmus of Tehuantepec in eastern Oaxaca. Geography The Sierra Madre del Sur joins with the Eje Volcánico Transv ...
mountains, from Guerrero to southern Oaxaca. "Red warbler" has been designated the official name by the IOC. It is a straightforward reference to its color. The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
name ''Cardellina'' is the diminutive of the Italian ''cardella'', a regional name for the European goldfinch, while its
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''rubra'', is
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for "red".


Description

The red warbler is a small passerine, measuring in length, and weighing from . As an adult, it is red overall, with either a white or dark gray (depending on the subspecies) auricular patch on each side of its head. Its wings and tail are slightly darker, dusky red, and edged in pinkish-red. Its legs are a dull red-brown, and its thin
bill Bill(s) may refer to: Common meanings * Banknote, paper cash (especially in the United States) * Bill (law), a proposed law put before a legislature * Invoice, commercial document issued by a seller to a buyer * Bill, a bird or animal's beak Plac ...
is pinkish-gray with a dark tip. The iris is dark brown to blackish. Plumage varies little between the sexes, although the female tends to be a little duller or more orange-tinged. Adult pairs separate and molt fully from August, after the breeding season. The adult red warbler is hard to confuse with any other bird species in its range; the scarlet tanager and
summer tanager The summer tanager (''Piranga rubra'') is a medium-sized American songbird. Formerly placed in the tanager family (Thraupidae), it and other members of its genus are now classified in the cardinal family (Cardinalidae). The species's plumage and ...
have similar mostly-red plumage but are larger with thick conical bills. As a juvenile, the red warbler is pinkish-brown with a whitish auricular patch. Its darker wings and tail show pinkish-cinnamon edges, with two paler on the former.


Voice

The red warbler has several common calls, including a high, thin ''tsii'' and a stronger ''pseet''. Its
song A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetitio ...
is a mix of short trills and richer warbles, interspersed with high-pitched chips. Unlike other species in the same
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
zone, it tends to sing only during bright morning hours during the
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
. It does not sing – and even its calling frequency decreases – in cloudy weather, regardless of season.


Distribution and habitat

Endemic to the highlands of Mexico north of the Isthmus of Tehuantepec, the red warbler has three disjunct populations that correspond to the three subspecies: from southwestern Chihuahua to northern Nayarit, from southern Jalisco and southern Hidalgo to Oaxaca, and from Guerrero into southern Oaxaca in the
Sierra Madre del Sur The Sierra Madre del Sur is a mountain range in southern Mexico, extending from southern Michoacán east through Guerrero, to the Isthmus of Tehuantepec in eastern Oaxaca. Geography The Sierra Madre del Sur joins with the Eje Volcánico Transv ...
mountains. It is fairly common to common in the country's interior and on adjacent slopes, where it occurs at elevations ranging from above sea level; it does not occur along either coast. It is an altitudinal migrant, moving from higher humid or semi-humid
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
, pine-
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
and
fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...
forests in the breeding season to lower elevations, often in oak forests, in the winter. It is among the most common of the small birds in its woodland habitat, second only to the
golden-crowned kinglet The golden-crowned kinglet (''Regulus satrapa'') is a very small songbird in the family Regulidae that lives throughout much of North America. Description Adults are olive-gray on the upperparts with white underparts, with thin bills and shor ...
in fir forests in one study and the third most common warbler in oak-conifer woodlands in another. Though the species was reported to have been collected in
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
in the late 19th century, the record's location was not widely believed, and there is no strong evidence that it ever occurred there. It has strayed as far north as southeastern Arizona, where a bird was found on Mount Lemmon in 2018.


Behavior

Though it occasionally joins
mixed-species flock A mixed-species feeding flock, also termed a mixed-species foraging flock, mixed hunting party or informally bird wave, is a flock of usually insectivorous birds of different species that join each other and move together while foraging. These ar ...
s, the red warbler is more typically found alone or in pairs. Youngsters probably choose mates in the autumn of their first year, and pairs remain together year-round, except during severe weather and during post-breeding molt.


Breeding

The red warbler breeds primarily in early spring, from February until May, though at least one nest containing young has been found as late as the end of June. By late February, the male establishes a territory that averages about 40 square metres, defending it with song. Other males may intrude by silently flying in at a height of around 3 m (10 ft) and back to their own territory. Deep incursions result in combat, after which the interloper usually leaves. Beginning in mid-March, the male courts the female by chasing her through the undergrowth. The pair then perch together while the male sings and the female calls softly. The female alone builds the nest, a task which typically takes 4–6 days. She chooses a sunlit area, such as an area of windfall, the brushy edge of a trail or water course, or a small clearing, for its location. Tightly woven of plant material, the nest is hidden in ground vegetation and anchored to the stalks of surrounding vegetation. The nest, which is usually
oven upA double oven A ceramic oven An oven is a tool which is used to expose materials to a hot environment. Ovens contain a hollow chamber and provide a means of heating the chamber in a controlled way. In use since antiquity, they have been us ...
-shaped with a side or upward-facing entrance, measures roughly wide by long by high. Bulky and untidy on the outside, it is typically constructed primarily of dead pine needles and dead grass, though gray
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship. shreds of bark and tips of fern fronds are also used; most of these materials are gathered from the ground close to the nest, though some is picked from low branches or further away. A few nests are only cups, lacking the roof of the more typical structures. Inside, the nest is tidy and compact, lined with fine grasses and plant fluff, which is generally gathered some distance from the nest. Early in the breeding season, there may be a gap of as many as 11 days between the completion of the nest and the laying of the first
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
. Later in the season, this time decreases so that the first egg is laid as soon as the nest is ready. The female normally lays three eggs, though
clutches A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...
of up to four have been recorded. The eggs, which are variously described as pale pink with evenly distributed brown spots or white with cinnamon and rust spots densely ringing the larger end of the egg, measure by and weigh . The female alone incubates the eggs for 16 days; the male does not even approach the nest until several days after the eggs hatch. She sits facing the back wall of the nest, with her head and body sheltered by its roof and her tail sticking out the opening. She sits tight at the approach of danger, typically not flying until a potential predator actually makes contact with the nest. Both adults feed the nestlings and remove
fecal sac A fecal sac (also spelled faecal sac) is a mucous membrane, generally white or clear with a dark end, that surrounds the feces of some species of nestling birds. It allows parent birds to more easily remove fecal material from the nest. The nes ...
s, though the female removes far more than the male does. The parents move deceptively when approaching the nest, foraging – or pretending to forage – in nearby vegetation. They stay only a few seconds in any one spot, including at the nest, making it more difficult for a predator to locate the young. Nestlings make a rapid, high-pitched peeping call as an adult approaches carrying food. They
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
within 10–11 days of hatching. Young birds are fully grown three weeks after fledging, upon which time they are driven off by their parents.


Food and feeding

The red warbler is an
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
. It
gleans Gleaning is the act of collecting leftover crops from farmers' fields after they have been commercially harvested or on fields where it is not economically profitable to harvest. It is a practice described in the Hebrew Bible that became a legall ...
primarily in understory shrubs at low to middle levels, moving slowly and deliberately through more open areas of the vegetation, and feeding with quick jabs into cracks in
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
and
pine needle A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
clusters. It sometimes hovers briefly to feed at pine needle clusters, a foraging technique known as "hover gleaning". Though it lacks any obvious adaptations for climbing, it regularly does so in its search for prey items on bark and
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s on branches, often hanging head-down as it probes. In areas of
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
growth, it typically flycatches, making brief aerial sorties from a perch in pursuit of flying insects. While it seldom associates with mixed-species flocks, it often feeds alongside other birds with no signs of conflict, displaying no hostility towards other species—such as the slate-throated whitestart (''Myioborus miniatus'')—with which it competes. It has been observed chasing off a flycatcher of the genus '' Empidonax''. Its foraging area is quite small, often amounting to only a few dozen square meters (several hundred square feet) per day. Late in the afternoon, its rate of foraging declines, and it rests, often taking brief naps, in the forest understory. Though it does not generally feed after sunset, it may do so to take advantage of transient food sources, such as hatching
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
.


Parasitism and predation

The red warbler is presumably hunted by small hawks such as the sharp-shinned hawk, and its nest raided by wrens, rodents, raccoons, feral cats and snakes. '' Isospora cardellinae'' is a
protozoan Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
species that has been isolated from a red warbler from Nevado de Toluca National Park, Mexico. It is a parasite that lives in cells in the villi of the bird's small intestine.


Toxicity

In the 16th century, Friar Bernardino de Sahagún had reported that a red bird matching the description of the red warbler was regarded as inedible by the Aztecs. Researchers Patricia Escalante and John W. Daly isolated two alkaloids in preliminary investigations of the feathers. The presence of these alkaloids render the bird unpalatable; humans find it inedible.


Conservation and threats

The red warbler is currently rated as a species of
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. Though there is evidence that its numbers are decreasing, the decline has not been precipitous (that is, less than a 30% decline over ten years or three generations), and the population remains quite large, estimates ranging from 50,000 to 499,999 birds. The forested areas in which it occurs include some of the most threatened habitats in Mexico; logging, agricultural expansion,
firewood Firewood is any wooden material that is gathered and used for fuel. Generally, firewood is not highly processed and is in some sort of recognizable log or branch form, compared to other forms of wood fuel like pellets or chips. Firewood can ...
gathering, road building, tourist development, overgrazing and intensive urbanization are among the many things contributing to the destruction of the forests. There is some evidence that selective logging in pine forests may actually favor this species, which prefers more open, sunlit areas in which to breed.


Note


References


External links

*
Red warbler photos
on the Academy of Natural Sciences' Visual Resources for Ornithology website
Red warbler videos
on the Internet Bird Collection website
Red warbler vocalizations
on the Macauley Library's (
Cornell Laboratory of Ornithology The Cornell Lab of Ornithology is a member-supported unit of Cornell University in Ithaca, New York, which studies birds and other wildlife. It is housed in the Imogene Powers Johnson Center for Birds and Biodiversity in Sapsucker Woods Sanctuar ...
) website
Red warbler vocalizations
on the xeno-canto.org website {{Authority control red warbler Birds of Mexico Endemic birds of Mexico red warbler Taxa named by William John Swainson Birds of the Sierra Madre Occidental Birds of the Sierra Madre del Sur Birds of the Trans-Mexican Volcanic Belt