Hummingbird

Hummingbirds are birds native to the Americas and comprise

the biological family Trochilidae. With about 360 species, they Hummingbird
occur from Alaska to Tierra del Fuego, but the vast majority of Temporal range: Rupelian
the species are found in the tropics. They are small birds, with
most species measuring 7.5–13  cm (3–5  in) in length. The
smallest extant hummingbird species is the 5  cm (2.0  in) bee
hummingbird, which weighs less than 2.0  g (0.07  oz). The
largest hummingbird species is the 23  cm (9.1  in) giant
hummingbird, weighing 18–24 grams (0.63–0.85 oz). They are
specialized for feeding on flower nectar, but all species also
consume flying insects or spiders.

Hummingbirds split from their sister group, the swifts and

treeswifts, around 42 million years ago. The common ancestor of
extant hummingbirds is estimated to have lived 22 million years
ago in South America. They are known as hummingbirds Four hummingbirds
because of the humming sound created by their beating wings,
from Trinidad and Tobago
which flap at high frequencies audible to humans. They hover in
mid-air at rapid wing-flapping rates, which vary from around 12 Scientific classification
beats per second in the largest species to around 80 per second in
Kingdom: Animalia
small hummingbirds. Of those species that have been measured
during flying in wind tunnels, their top speeds exceed 15  m/s Phylum: Chordata
(54  km/h; 34  mph). During courtship, some male species dive
from 30 metres (100  ft) of height above a female at speeds Class: Aves
around 23 m/s (83 km/h; 51 mph).[1][2] Order: Apodiformes

Hummingbirds have the highest mass-specific metabolic rate of Family: Trochilidae

any homeothermic animal.[3] To conserve energy when food is Vigors, 1825
scarce and at night when not foraging, they can enter torpor, a
Subfamilies
state similar to hibernation, and slow their metabolic rate to 1/15
of its normal rate.[4]
†Eurotrochilus
Florisuginae
Phaethornithinae
Contents Polytminae
Lesbiinae
Taxonomy and systematics Patagoninae
Evolution Trochilinae
Sexual dimorphisms For an alphabetic species list,
Co-evolution with ornithophilous flowers see List of hummingbird species
Bill evolution
Specialized characteristics and metabolism
Humming
Wingbeats and flight stability
Vision
 Metabolism
Heat dissipation
Kidney function
Song and vocal learning
Torpor
Lifespan
Predators
Reproduction
Feather colors
Aerodynamics of flight
Feather sonation
Courtship dives
Wing feather trill
Range
Migration
Diet and specializations for food gathering
Perception of sweet nectar
Tongue as a micropump
Feeders and artificial nectar
Visual cues of foraging
Olfaction
Superficially similar species
In myth and culture
Gallery
See also
References
External links

Taxonomy and systematics

The family Trochilidae was introduced in 1825 by Irish zoologist Nicholas Aylward Vigors with Trochilus
as the type genus.[5][6] Molecular phylogenetic studies of the hummingbirds have shown that the family is
composed of nine major clades.[7][8] When Edward Dickinson and James Van Remsen Jr. updated the
Howard and Moore Complete Checklist of the Birds of the World for the 4th edition in 2013, they divided
the hummingbirds into six subfamilies.[9]

Trochilidae
   
  Florisuginae – topazes
 
 
 
Phaethornithinae – hermits
 
 
 
  Polytminae – mangoes
 
   Lesbiinae   Heliantheini – brilliants
     
 
  Lesbiini – coquettes

 
  Patagoninae – giant hummingbird

 
Lampornithini – mountain gems
 
   
     
Trochilinae Mellisugini – bees
   

 
  Trochilini – emeralds

The above cladogram of the hummingbird family is based on a molecular phylogenetic study by Jimmy
McGuire and collaborators published in 2014.[8] The English names are those introduced by Robert
Bleiweiss, John Kirsch, and Juan Matheus in 1997.[10] The Latin names are those proposed by Dickinson
and Remsen in 2013.[9]

In traditional taxonomy, hummingbirds are placed in the order

Apodiformes, which also contains the swifts, but some taxonomists
have separated them into their own order, the Trochiliformes.
Hummingbirds' wing bones are hollow and fragile, making
fossilization difficult and leaving their evolutionary history poorly
documented. Though scientists theorize that hummingbirds
originated in South America, where species diversity is greatest,
possible ancestors of extant hummingbirds may have lived in parts
of Europe and what is southern Russia today.[11]

Around 360 hummingbirds have been described. They have been

traditionally divided into two subfamilies: the hermits (subfamily
Phaethornithinae) and the typical hummingbirds (subfamily
Trochilinae, all the others). Molecular phylogenetic studies have
shown, though, that the hermits are sister to the topazes, making the
former definition of the Trochilinae not monophyletic. The
hummingbirds form nine major clades: the topazes and jacobins,
the hermits, the mangoes, the coquettes, the brilliants, the giant A color plate illustration from Ernst
hummingbird (Patagona gigas), the mountaingems, the bees, and Haeckel's Kunstformen der Natur
the emeralds.[8] The topazes and jacobins combined have the oldest (1899), showing a variety of
split with the rest of the hummingbirds. The hummingbird family hummingbirds
has the third-greatest number of species of any bird family (after the
tyrant flycatchers and the tanagers).[8][12]

Fossil hummingbirds are known from the Pleistocene of Brazil and the Bahamas, but neither has yet been
scientifically described, and fossils and subfossils of a few extant species are known. Until recently, older
fossils had not been securely identifiable as those of hummingbirds.

In 2004, Gerald Mayr identified two 30-million-year-old hummingbird fossils. The fossils of this primitive
hummingbird species, named Eurotrochilus inexpectatus ("unexpected European hummingbird"), had been
sitting in a museum drawer in Stuttgart; they had been unearthed in a clay pit at Wiesloch–Frauenweiler,
south of Heidelberg, Germany, and, because hummingbirds were assumed to have never occurred outside
the Americas, were not recognized to be hummingbirds until Mayr took a closer look at them.[11][13]

Fossils of birds not clearly assignable to either hummingbirds or a related extinct family, the Jungornithidae,
have been found at the Messel pit and in the Caucasus, dating from 35 to 40 million years ago; this
indicates that the split between these two lineages indeed occurred around that time. The areas where these
early fossils have been found had a climate quite similar to that of the northern Caribbean or southernmost
China during that time. The biggest remaining mystery at present is what happened to hummingbirds in the
roughly 25 million years between the primitive Eurotrochilus and the modern fossils. The astounding
morphological adaptations, the decrease in size, and the dispersal to the Americas and extinction in Eurasia
all occurred during this timespan. DNA–DNA hybridization results suggest that the main radiation of South
American hummingbirds took place at least partly in the Miocene, some 12 to 13 million years ago, during
the uplifting of the northern Andes.[14]

In 2013, a 50-million-year-old bird fossil unearthed in Wyoming was found to be a predecessor to both
hummingbirds and swifts before the groups diverged.[15]

Evolution
Hummingbirds are thought to have split from other members of Apodiformes, the insectivorous swifts
(family Apodidae) and treeswifts (family Hemiprocnidae), about 42 million years ago, probably in
Eurasia.[8] Despite their current New World distribution, the earliest known species of hummingbird are
known from the early Oligocene (Rupelian ~34–28 million years ago) of Europe, belonging to the genus
Eurotrochilus, which is very similar in its morphology to modern hummingbirds.[13][16][17] A phylogenetic
tree unequivocally indicates that modern hummingbirds originated in South America, with the last common
ancestor of all living hummingbirds living around 22 million years ago.[8]

A map of the hummingbird family tree  – reconstructed from analysis of 284 of the world's 338 known
species  – shows rapid diversification from 22 million years ago.[18] Hummingbirds fall into nine main
clades – the topazes, hermits, mangoes, brilliants, coquettes, the giant hummingbird, mountaingems, bees,
and emeralds – defining their relationship to nectar-bearing flowering plants and the birds' continued spread
into new geographic areas.[18][7][8][19]

While all hummingbirds depend on flower nectar to fuel their high metabolisms and hovering flight,
coordinated changes in flower and bill shape stimulated the formation of new species of hummingbirds and
plants. Due to this exceptional evolutionary pattern, as many as 140 hummingbird species can coexist in a
specific region, such as the Andes range.[18]

The hummingbird evolutionary tree shows one key evolutionary factor appears to have been an altered
taste receptor that enabled hummingbirds to seek nectar.[20]

The Andes Mountains appear to be a particularly rich environment for hummingbird evolution because
diversification occurred simultaneously with mountain uplift over the past 10 million years.[18]
Hummingbirds remain in dynamic diversification inhabiting ecological regions across South America,
North America, and the Caribbean, indicating an enlarging evolutionary radiation.[18]

Within the same geographic region, hummingbird clades co-evolved with nectar-bearing plant clades,
affecting mechanisms of pollination.[21][22] The same is true for the sword-billed hummingbird (Ensifera
ensifera), one of the morphologically most extreme species, and one of its main food plant clades
(Passiflora section Tacsonia).[23]
Sexual dimorphisms

Hummingbirds exhibit sexual size dimorphism according to Rensch's rule,[24]

in which males are smaller than females in small-bodied species, and males are
larger than females in large-bodied species.[25] The extent of this sexual size
difference varies among clades of hummingbirds.[25][26] For example, the
Mellisugini clade (bees) exhibits a large size dimorphism, with females being
larger than males.[26] Conversely, the Lesbiini clade (coquettes) displays very
little size dimorphism; males and females are similar in size.[26]
Sexual
dimorphisms in bill size and shape are also present between male and female
hummingbirds,[26] where in many clades, females have longer, more curved
bills favored for accessing nectar from tall flowers.[27] For males and females
of the same size, females tend to have larger bills.[26] Violet-tailed sylph resting
on a branch in
Sexual size and bill differences likely evolved due to constraints imposed by northwestern Ecuador
courtship, because mating displays of male hummingbirds require complex
aerial maneuvers.[24] Males tend to be smaller than females, allowing
conservation of energy to forage competitively and participate more frequently
in courtship.[24] Thus, sexual selection favors smaller male hummingbirds.[24]

Female hummingbirds tend to be larger, requiring more energy, with longer

beaks that allow for more effective reach into crevices of tall flowers for
nectar.[27] Thus, females are better at foraging, acquiring flower nectar, and
supporting the energy demands of their larger body size.[27] Directional
selection thus favors the larger hummingbirds in terms of acquiring food.[25]

Another evolutionary cause of this sexual bill dimorphism is that the selective
forces from competition for nectar between the sexes of each species drives Sexual dimorphism
sexual dimorphism.[26] Depending on which sex holds territory in the species, displayed in violet-tailed
the other sex having a longer bill and being able to feed on a wide variety of sylph male (top) and
flowers is advantageous, decreasing intraspecific competition.[27] For female (bottom)
example, in species of hummingbirds where males have longer bills, males do
not hold a specific territory and have a lek mating system.[27] In species where
males have shorter bills than females, males defend their resources, so females benefit from a longer bill to
feed from a broader range of flowers.[27]

Co-evolution with ornithophilous flowers

Hummingbirds are specialized nectarivores[28] and are tied to the

ornithophilous flowers upon which they feed. This coevolution
implies that morphological traits of hummingbirds, such as bill
length, bill curvature, and body mass are correlated with
morphological traits of plants, for example corolla length,
curvature, and volume.[29] Some species, especially those with
unusual bill shapes, such as the sword-billed hummingbird and the
sicklebills, are co-evolved with a small number of flower species. Purple-throated carib feeding at a
Even in the most specialized hummingbird–plant mutualisms, flower
though, the number of food plant lineages of the individual
hummingbird species increases with time.[30] The bee
hummingbird (Mellisuga helenae) – the world's smallest bird – evolved to dwarfism likely because it had to
compete with long-billed hummingbirds having an advantage for nectar foraging from specialized flowers,
consequently leading the bee hummingbird to more successfully compete for flower foraging against
insects.[31][32]

Many plants pollinated by hummingbirds produce flowers in shades of red, orange, and bright pink, though
the birds take nectar from flowers of other colors as well. Hummingbirds can see wavelengths into the near-
ultraviolet, but hummingbird-pollinated flowers do not reflect these wavelengths as many insect-pollinated
flowers do. This narrow color spectrum may render hummingbird-pollinated flowers relatively
inconspicuous to most insects, thereby reducing nectar robbing.[33][34] Hummingbird-pollinated flowers
also produce relatively weak nectar (averaging 25% sugars w/w) containing a high proportion of sucrose,
whereas insect-pollinated flowers typically produce more concentrated nectars dominated by fructose and
glucose.[35]

Hummingbirds and the plants they visit for nectar have a tight co-evolutionary association, generally called
a plant–bird mutualistic network.[36] These birds show high specialization and modularity, especially in
communities with high species richness. These associations are also observed when closely related
hummingbirds, for example two species of the same genus, visit distinct sets of flowering species.[36][37]

Bill evolution

Upon maturity, males of a particular species, Phaethornis longirostris, the long-billed hermit, appear to be
evolving a dagger-like weapon on the beak tip as a secondary sexual trait to defend mating areas.[38]

Specialized characteristics and metabolism

Humming

Hummingbirds are named for the prominent humming sound their

0:00 / 0:00
wingbeats make while flying and hovering to feed or interact with
other hummingbirds.[39] Humming serves communication purposes
by alerting other birds of the arrival of a fellow forager or potential A calliope hummingbird hovering
near a feeder, creating the
mate.[39] The humming sound derives from aerodynamic forces
"humming" sound from its rapid
generated by both the downstrokes and upstrokes of the rapid
wingbeats, while chirping by
wingbeats, causing oscillations and harmonics that evoke an
vocalization
acoustic quality likened to that of a musical instrument.[39][40] The
humming sound of hummingbirds is unique among flying animals,
compared to the whine of mosquitoes, buzz of bees, and "whoosh" of larger birds.[39][40]

The wingbeats causing the hum of hummingbirds during hovering are achieved by elastic recoil of wing
strokes produced by the main flight muscles: the pectoralis major (the main downstroke muscle) and
supracoracoideus (the main upstroke muscle).[41]

Wingbeats and flight stability

The highest recorded wingbeats for wild hummingbirds during hovering is 88/sec, as measured for the
purple-throated woodstar (Calliphlox mitchellii) weighing 3.2  g.[42] The number of beats per second
increases above "normal" while hovering during courtship displays (up to 90/sec for the calliope
hummingbird, Stellula calliope), a wingbeat rate 40% higher than its typical hovering rate.[43]
During turbulent airflow conditions created experimentally in a wind tunnel, hummingbirds exhibit stable
head positions and orientation when they hover at a feeder.[44] When wind gusts from the side,
hummingbirds compensate by increasing wing-stroke amplitude and stroke plane angle and by varying
these parameters asymmetrically between the wings and from one stroke to the next.[44] They also vary the
orientation and enlarge the collective surface area of their tail feathers into the shape of a fan.[44] While
hovering, the visual system of a hummingbird is able to separate apparent motion caused by the movement
of the hummingbird itself from motions caused by external sources, such as an approaching predator.[45] In
natural settings full of highly complex background motion, hummingbirds are able to precisely hover in
place by rapid coordination of vision with body position.[45]

Vision

Although hummingbird eyes are small in diameter (5–6  mm), they are accommodated in the skull by
reduced skull ossification, and occupy a relatively larger proportion of the skull compared to other birds and
animals.[46] Further, hummingbird eyes have relatively large corneas, which comprise about 50% of the
total transverse eye diameter, combined with an extraordinary density of retinal ganglion cells responsible
for visual processing, containing some 45,000 neurons per mm2 .[47] The enlarged cornea relative to total
eye diameter serves to increase the amount of light perception by the eye when the pupil is dilated
maximally, enabling nocturnal flight.[47]

During evolution, hummingbirds adapted to the navigational needs of visual processing while in rapid flight
or hovering by development of the exceptionally dense array of retinal neurons, allowing for increased
spatial resolution in the lateral and frontal visual fields.[47] Morphological studies of the hummingbird brain
showed that neuronal hypertrophy – relatively the largest in any bird – exists in a region called the pretectal
nucleus lentiformis mesencephali (called the nucleus of the optic tract in mammals) responsible for refining
dynamic visual processing while hovering and during rapid flight.[48][49]

The enlargement of the brain region responsible for visual processing indicates an enhanced ability for
perception and processing of fast-moving visual stimuli which hummingbirds encounter during rapid
forward flight, insect foraging, competitive interactions, and high-speed courtship.[49][50] A study of broad-
tailed hummingbirds indicated that hummingbirds have a fourth color-sensitive visual cone (humans have
three) that detects ultraviolet light and enables discrimination of non-spectral colors, possibly having a role
in courtship displays, territorial defense, and predator evasion.[51] The fourth color cone would extend the
range of visible colors for hummingbirds to perceive ultraviolet light and color combinations of feathers and
gorgets, colorful plants, and other objects in their environment, enabling detection of as many as five non-
spectral colors, including purple, ultraviolet-red, ultraviolet-green, ultraviolet-yellow, and ultraviolet-
purple.[51]

Hummingbirds are highly sensitive to stimuli in their visual fields, responding to even minimal motion in
any direction by reorienting themselves in midflight.[45][49][50] Their visual sensitivity allows them to
precisely hover in place while in complex and dynamic natural environments,[45] functions enabled by the
lentiform nucleus which is tuned to fast-pattern velocities, enabling highly tuned control and collision
avoidance during forward flight.[49]

Metabolism

With the exception of insects, hummingbirds while in flight have the highest metabolism of all animals – a
necessity to support the rapid beating of their wings during hovering and fast forward flight.[3][52] Their
heart rate can reach as high as 1,260 bpm, a rate once measured in a blue-throated hummingbird, with a
breathing rate of 250 bpm, even at rest.[53][54] During flight, oxygen consumption per gram of muscle
tissue in a hummingbird is about 10 times higher than that measured in elite human athletes.[3]

Hummingbirds are rare among vertebrates in their ability to rapidly make use of ingested sugars to fuel
energetically expensive hovering flight,[55] powering up to 100% of their metabolic needs with the sugars
they drink (in comparison, human athletes maximize around 30%). Hummingbirds can use newly ingested
sugars to fuel hovering flight within 30–45 minutes of consumption.[56][57] These data suggest that
hummingbirds are able to oxidize sugar in flight muscles at rates high enough to satisfy their extreme
metabolic demands. A 2017 review indicated that hummingbirds have in their flight muscles a mechanism
for "direct oxidation" of sugars into maximal ATP yield to support their high metabolic rate for hovering,
foraging at altitude, and migrating.[58]

By relying on newly ingested sugars to fuel flight, hummingbirds can reserve their limited fat stores to
sustain their overnight fasting or to power migratory flights.[56] Studies of hummingbird metabolism
address how a migrating ruby-throated hummingbird can cross 800 km (500 mi) of the Gulf of Mexico on a
nonstop flight.[54] This hummingbird, like other long-distance migrating birds, stores fat as a fuel reserve
augmenting its weight by as much as 100%, then enabling metabolic fuel for flying over open water.[54][59]

Heat dissipation

The high metabolic rate of hummingbirds – especially during rapid forward flight and hovering – produces
increased body heat that requires specialized mechanisms of thermoregulation for heat dissipation, which
becomes an even greater challenge in hot, humid climates.[60] Hummingbirds dissipate heat partially by
evaporation through exhaled air, and from body structures with thin or no feather covering, such as around
the eyes, shoulders, under the wings (patagia), and feet.[61][62]

While hovering, hummingbirds do not benefit from the heat loss by air convection during forward flight,
except for air movement generated by their rapid wing-beat, possibly aiding convective heat loss from the
extended feet.[60][63] Smaller hummingbird species, such as the calliope, appear to adapt their relatively
higher surface-to-volume ratio to improve convective cooling from air movement by the wings.[60] When
air temperatures rise above 36  °C (97  °F), thermal gradients driving heat passively by convective
dissipation from around the eyes, shoulders, and feet are reduced or eliminated, requiring heat dissipation
mainly by evaporation and exhalation.[60] In cold climates, hummingbirds retract their feet into breast
feathers to eliminate skin exposure and minimize heat dissipation.[63]

Kidney function

The dynamic range of metabolic rates in hummingbirds[64] requires a parallel dynamic range in kidney
function.[65] During a day of nectar consumption with a corresponding high water intake that may total five
times the body weight per day, hummingbird kidneys process water via glomerular filtration rates (GFR) in
amounts proportional to water consumption, thereby avoiding overhydration.[65][66] During brief periods of
water deprivation, however, such as in nighttime torpor, GFR drops to zero, preserving body water.[65][66]

Hummingbird kidneys also have a unique ability to control the levels of electrolytes after consuming
nectars with high amounts of sodium and chloride or none, indicating that kidney and glomerular structures
must be highly specialized for variations in nectar mineral quality.[67] Morphological studies on Anna's
hummingbird kidneys showed adaptations of high capillary density in close proximity to nephrons,
allowing for precise regulation of water and electrolytes.[66][68]
Song and vocal learning

Consisting of chirps, squeaks, whistles and buzzes,[69] hummingbird songs originate from at least seven
specialized nuclei in the forebrain.[70][71] A genetic expression study showed that these nuclei enable vocal
learning (ability to acquire vocalizations through imitation), a rare trait known to occur in only two other
groups of birds (parrots and songbirds) and a few groups of mammals (including humans, whales and
dolphins, and bats).[70] Within the past 66 million years, only hummingbirds, parrots, and songbirds out of
23 bird orders may have independently evolved seven similar forebrain structures for singing and vocal
learning, indicating that evolution of these structures is under strong epigenetic constraints possibly derived
from a common ancestor.[70][72]

The blue-throated hummingbird's song differs from typical oscine songs in its wide frequency range,
extending from 1.8  kHz to about 30  kHz.[73] It also produces ultrasonic vocalizations which do not
function in communication.[73] As blue-throated hummingbirds often alternate singing with catching small
flying insects, it is possible the ultrasonic clicks produced during singing disrupt insect flight patterns,
making insects more vulnerable to predation.[73]

The avian vocal organ, the syrinx, plays an important role in understanding hummingbird song
production.[74] What makes the hummingbird's syrinx different from that of other birds in the Apodiformes
order is the presence of internal muscle structure, accessory cartilages, and a large tympanum that serves as
an attachment point for external muscles, all of which are adaptations thought to be responsible for the
hummingbird's increased ability in pitch control and large frequency range.[74][75]

Torpor

The metabolism of hummingbirds can slow at night or at any time when food is not readily available; the
birds enter a hibernatory, deep-sleep state (known as torpor) to prevent energy reserves from falling to a
critical level. During nighttime torpor, body temperature falls from 40 to 18 °C,[76] with heart and breathing
rates both slowed dramatically (heart rate of roughly 50 to 180 bpm from its daytime rate of higher than
1000 bpm).[77]

During torpor, to prevent dehydration, the GFR ceases, preserving needed compounds such as glucose,
water, and nutrients.[65] Further, body mass declines throughout nocturnal torpor at a rate of 0.04 g per
hour, amounting to about 10% of weight loss each night.[65] The circulating hormone, corticosterone, is
one signal that arouses a hummingbird from torpor.[78]

Use and duration of torpor vary among hummingbird species and are affected by whether a dominant bird
defends territory, with nonterritorial subordinate birds having longer periods of torpor.[79] The
hummingbirds of the Andes in South America are known for entering exceptionally deep torpor and
dropping their body temperature.[80]

Lifespan

Hummingbirds have unusually long lifespans for organisms with such rapid metabolisms. Though many
die during their first year of life, especially in the vulnerable period between hatching and fledging, those
that survive may occasionally live a decade or more.[81] Among the better-known North American species,
the typical lifespan is probably 3 to 5 years.[81] For comparison, the smaller shrews, among the smallest of
all mammals, seldom live longer than 2 years.[82] The longest recorded lifespan in the wild relates to a
female broad-tailed hummingbird that was banded (ringed) as an adult at least one year old, then recaptured
11 years later, making her at least 12 years old.[83] Other longevity records for banded hummingbirds
include an estimated minimum age of 10 years 1 month for a female black-chinned hummingbird similar in
size to the broad-tailed hummingbird, and at least 11 years 2 months for a much larger buff-bellied
hummingbird.[84]

Predators

Praying mantises have been observed as predators of hummingbirds.[85][86][87] Other predators include
dragonflies, frogs, orb-weaver spiders, and other birds, such as the roadrunner.[88]

Reproduction
As far as is known, male hummingbirds do not take part in nesting.[89] Most species build a cup-shaped
nest on the branch of a tree or shrub,[90] although a few tropical species normally attach their nests to
leaves. The nest varies in size relative to the particular species – from smaller than half a walnut shell to
several centimeters in diameter.[89][91]

Many hummingbird species use spider silk and lichen to bind the nest material together and secure the
structure.[90][92] The unique properties of the silk allow the nest to expand as the young hummingbirds
grow.[90][91] Two white eggs are laid,[90][92] which despite being the smallest of all bird eggs are large
relative to the adult hummingbird's size.[91] Incubation lasts 14 to 23 days,[92] depending on the species,
ambient temperature, and female attentiveness to the nest.[89] The mother feeds her nestlings on small
arthropods and nectar by inserting her bill into the open mouth of a nestling, and then regurgitating the food
into its crop.[89][91] Hummingbirds stay in the nest for 18–22 days, after which they leave the nest to forage
on their own, although the mother bird may continue feeding them for another 25 days.[93]


Play media
Incubating in Copiapó, Chile Nest with two
Hummingbird building eggs in San
a nest in San Diego Jose, California
Zoo, video

Nest with two nestlings Feeding two nestlings in Fallen Anna's

in Santa Monica, Grand Teton National Park hummingbird nest in
California Ventura County,
California, shown next
to a toothpick for scale

Feather colors
To serve courtship and territorial competition, many male hummingbirds
have plumage with bright, varied coloration[94] resulting both from
pigmentation in the feathers and from prismal cells within the top layers of
feathers of the head, gorget, breast, back and wings.[95] When sunlight hits
these cells, it is split into wavelengths that reflect to the observer in varying
degrees of intensity,[95] with the feather structure acting as a diffraction
grating.[95] Iridescent hummingbird colors result from a combination of
refraction and pigmentation, since the diffraction structures themselves are
made of melanin, a pigment,[94] and may also be colored by carotenoid
pigmentation and more subdued black, brown or gray colors dependent on
melanin.[95]
Male Anna's hummingbird
By merely shifting position, feather regions of a muted-looking bird can showing iridescent crown
instantly become fiery red or vivid green.[95] In courtship displays for one and gorget feathers
example, males of the colorful Anna's hummingbird orient their bodies and
feathers toward the sun to enhance the display value of iridescent plumage
toward a female of interest.[96]

One study of Anna's hummingbirds found that dietary protein was an influential factor in feather color, as
birds receiving more protein grew significantly more colorful crown feathers than those fed a low-protein
diet.[97] Additionally, birds on a high-protein diet grew yellower (higher hue) green tail feathers than birds
on a low-protein diet.[97]

Aerodynamics of flight
Hummingbird flight has
been studied intensively
from an aerodynamic
perspective using wind
tunnels and high-speed
video cameras.

Two studies of rufous or

Anna's hummingbirds in a
A female ruby-throated hummingbird wind tunnel used particle
hovering in mid-air image velocimetry
techniques to investigate
the lift generated on the
bird's upstroke and downstroke. [99][100] The birds produced 75%
of their weight support during the downstroke and 25% during the
upstroke, with the wings making a "figure 8" motion.[101]

Many earlier studies had assumed that lift was generated equally A trail of wake vortices generated by
during the two phases of the wingbeat cycle, as is the case of a hummingbird's flight discovered
after training a bird to fly through a
insects of a similar size.[99] This finding shows that hummingbird
cloud of neutrally buoyant, helium-
hovering is similar to, but distinct from, that of hovering insects
filled soap bubbles and recording
such as the hawk moth.[99] Further studies using electromyography
airflows in the wake with stereo
in hovering rufous hummingbirds showed that muscle strain in the
photography.[98]
pectoralis major (principal downstroke muscle) was the lowest yet
recorded in a flying bird, and the primary upstroke muscle
(supracoracoideus) is proportionately larger than in other bird
species.[102]

Because of their flying technique, these birds no longer have an

alula.[103]

The giant hummingbird's wings beat as few as 12/sec[104] and the

wings of typical hummingbirds beat up to 80/sec.[105] Play media
Slow-motion video of hummingbirds
As air density decreases, for example, at higher altitudes, the feeding
amount of power a hummingbird must use to hover increases.
Hummingbird species adapted for life at higher altitudes, therefore,
have larger wings to help offset these negative effects of low air density on lift generation.[106]

A slow-motion video has shown how the hummingbirds deal with rain when they are flying. To remove
the water from their heads, they shake their heads and bodies, similar to a dog shaking, to shed water.[107]
Further, when raindrops collectively may weigh as much as 38% of the bird's body weight, hummingbirds
shift their bodies and tails horizontally, beat their wings faster, and reduce their wings' angle of motion
when flying in heavy rain.[108]
 High-speed capture sequence of two velvet-purple coronets sparring near a hummingbird
feeder in Ecuador

Feather sonation

Courtship dives

When courting, the male Anna's hummingbird ascends some 35 m (115 ft) above a female, before diving at
a speed of 27 m/s (89 ft/s), equal to 385 body lengths/sec – producing a high-pitched sound near the female
at the nadir of the dive.[109] This downward acceleration during a dive is the highest reported for any
vertebrate undergoing a voluntary aerial maneuver; in addition to acceleration, the speed, relative to body
length, is the highest known for any vertebrate. For instance, it is about twice the diving speed of peregrine
falcons in pursuit of prey.[109] At maximum descent speed, about 10 g of gravitational force occur in the
courting hummingbird during a dive (Note: G-force is generated as the bird pulls out of the dive).[109] By
comparison to humans, this is a G-force acceleration well beyond the threshold of causing near loss of
consciousness in fighter pilots (occurring at about +5 Gz) during flight of fixed-wing aircraft in a high-
speed banked turn.[109][110]

The outer tail feathers of male Anna's (Calypte anna) and Selasphorus hummingbirds (e.g., Allen's,
calliope) vibrate during courtship display dives and produce an audible chirp caused by aeroelastic
flutter.[111][112] Hummingbirds cannot make the courtship dive sound when missing their outer tail
feathers, and those same feathers could produce the dive sound in a wind tunnel.[111] The bird can sing at
the same frequency as the tail-feather chirp, but its small syrinx is not capable of the same volume.[113] The
sound is caused by the aerodynamics of rapid air flow past tail feathers, causing them to flutter in a
vibration, which produces the high-pitched sound of a courtship dive.[111][114]

Many other species of hummingbirds also produce sounds with their wings or tails while flying, hovering,
or diving, including the wings of the calliope hummingbird,[115] broad-tailed hummingbird, rufous
hummingbird, Allen's hummingbird, and streamertail, as well as the tail of the Costa's hummingbird and the
black-chinned hummingbird, and a number of related species.[116] The harmonics of sounds during
courtship dives vary across species of hummingbirds.[112]

Wing feather trill

Male rufous and broad-tailed hummingbirds (genus Selasphorus) have a distinctive wing feature during
normal flight that sounds like jingling or a buzzing shrill whistle.[117] The trill arises from air rushing
through slots created by the tapered tips of the ninth and tenth primary wing feathers, creating a sound loud
enough to be detected by female or competitive male hummingbirds and researchers up to 100  m
away.[117]

Behaviorally, the trill serves several purposes:[117]

 Announces the sex and presence of a male bird
Provides audible aggressive defense of a feeding territory and an intrusion tactic
Enhances communication of a threat
Favors mate attraction and courtship

Range
Hummingbirds are restricted to the Americas from south central Alaska to Tierra del Fuego, including the
Caribbean. The majority of species occur in tropical and subtropical Central and South America, but
several species also breed in temperate climates and some hillstars occur even in alpine Andean highlands
at altitudes up to 5,200 m (17,100 ft).[118]

The greatest species richness is in humid tropical and subtropical forests of the northern Andes and adjacent
foothills, but the number of species found in the Atlantic Forest, Central America or southern Mexico also
far exceeds the number found in southern South America, the Caribbean islands, the United States, and
Canada. While fewer than 25 different species of hummingbirds have been recorded from the United States
and fewer than 10 from Canada and Chile each,[119] Colombia alone has more than 160[120] and the
comparably small Ecuador has about 130 species.[121]

The migratory ruby-throated hummingbird breeds in a range from the Southeastern United States to
Ontario,[122] while the black-chinned hummingbird, its close relative and another migrant, is the most
widespread and common species in the southwestern United States. The rufous hummingbird is the most
widespread species in western North America,[123] and the only hummingbird to be recorded outside of the
Americas, having occurred in the Chukchi Peninsula of Russia.[124]

Migration
Most North American hummingbirds migrate southward in fall to spend winter in Mexico, the Caribbean
Islands, or Central America.[125] A few southern South American species also move north to the tropics
during the southern winter. A few species are year-round residents of Florida, California, and the far
southwestern desert regions of the US.[125] Among these are Anna's hummingbird, a common resident
from southern Arizona and inland California, and the buff-bellied hummingbird, a winter resident from
Florida across the Gulf Coast to South Texas. Ruby-throated hummingbirds are common along the Atlantic
flyway, and migrate in summer from as far north as Atlantic Canada,[125] returning to Mexico, South
America, southern Texas, and Florida to winter.[125][126] During winter in southern Louisiana, black-
chinned, buff-bellied, calliope, Allen's, Anna's, ruby-throated, rufous, broad-tailed, and broad-billed
hummingbirds are present.[125]

The rufous hummingbird breeds farther north than any other species of hummingbird,[125] often breeding
in large numbers in temperate North America and wintering in increasing numbers along the coasts of the
subtropical Gulf of Mexico and Florida, rather than in western or central Mexico.[127] By migrating in
spring as far north as the Yukon or southern Alaska,[125][127] the rufous hummingbird migrates more
extensively and nests farther north than any other hummingbird species, and must tolerate occasional
temperatures below freezing in its breeding territory. This cold hardiness enables it to survive temperatures
below freezing, provided that adequate shelter and food are available.[127]

As calculated by displacement of body size, the rufous hummingbird makes perhaps the longest migratory
journey of any bird in the world. At just over 3 in long, rufous birds travel 3,900 miles one-way from
Alaska to Mexico in late summer, a distance equal to 78,470,000 body lengths.[127] By comparison, the 13-
inch-long Arctic tern makes a one-way flight of about 11,185 miles, or 51,430,000 body lengths, just 65%
of the body displacement during migration by rufous hummingbirds.[127]

The northward migration of rufous hummingbirds occurs along the Pacific flyway[127] and may be time-
coordinated with flower and tree-leaf emergence in spring in early March, and also with availability of
insects as food.[128] Arrival at breeding grounds before nectar availability from mature flowers may
jeopardize breeding opportunities.[129]

Diet and specializations for food gathering

For nutrition,
hummingbirds eat a variety
of insects, including
mosquitoes, fruit flies, gnats
in flight, or aphids on
leaves and spiders in their
webs.[130][131][132][133] Play media
Lesser violetear at a flower The lower beak of Hummingbird in Copiapó, Chile: The
hummingbirds is flexible apparent slow movement of its wings
and can bend as much as is a result of the stroboscopic effect.
25 degrees when it widens at the base, making a larger surface for
catching insects.[132] Hummingbirds hover within insect swarms in
a method called "hover-hawking" to facilitate feeding.[133]

To supply energy needs, hummingbirds drink nectar, a sweet liquid inside certain flowers. Like bees, they
are able to assess the amount of sugar in the nectar they drink; they normally reject flower types that
produce nectar that is less than 10% sugar and prefer those whose sugar content is higher. Nectar is a
mixture of glucose, fructose, and sucrose, and is a poor source of other nutrients, requiring hummingbirds
to meet their nutritional needs by consuming insects.[132][133]

Hummingbirds do not spend all day flying, as the energy cost would be prohibitive; the majority of their
activity consists simply of sitting or perching. Hummingbirds eat many small meals and consume around
half their weight in nectar (twice their weight in nectar, if the nectar is 25% sugar) each day.[134]
Hummingbirds digest their food rapidly due to their small size and high metabolism; a mean retention time
less than an hour has been reported.[135] Hummingbirds spend an average of 10–15% of their time feeding
and 75–80% sitting and digesting.

Because their high metabolism makes them vulnerable to starvation, hummingbirds are highly attuned to
food sources. Some species, including many found in North America, are territorial and try to guard food
sources (such as a feeder) against other hummingbirds, attempting to ensure a future food supply for itself.
Additionally, hummingbirds have an enlarged hippocampus, a brain region facilitating spatial memory used
to map flowers previously visited during nectar foraging.[136]

Hummingbird beaks are flexible[132] and their shapes vary dramatically as an adaptation for specialized
feeding.[26][27] Some species, such as hermits (Phaethornis spp.) have long bills that allow them to probe
deep into flowers with long corollae. Thornbills have short, sharp bills adapted for feeding from flowers
with short corollae and piercing the bases of longer ones. The sicklebills' extremely decurved bills are
adapted to extracting nectar from the curved corollae of flowers in the family Gesneriaceae. The bill of the
fiery-tailed awlbill has an upturned tip, as in the avocets. The male tooth-billed hummingbird has
barracuda-like spikes at the tip of its long, straight bill.
The two halves of a hummingbird's bill have a pronounced overlap, with the lower half (mandible) fitting
tightly inside the upper half (maxilla). When a hummingbird feeds on nectar, the bill is usually opened only
slightly, allowing the tongue to dart out and into the interior of flowers. Hummingbird bill sizes range from
about 5 mm to as long as 100 mm (about 4 in).[137] When catching insects in flight, a hummingbird's jaw
flexes downward to widen the gape for successful capture.[131]

Perception of sweet nectar

Perception of sweetness in nectar evolved in hummingbirds during their genetic divergence from
insectivorous swifts, their closest bird relatives.[138] Although the only known sweet sensory receptor,
called T1R2,[139] is absent in birds, receptor expression studies showed that hummingbirds adapted a
carbohydrate receptor from the T1R1-T1R3 receptor, identical to the one perceived as umami in humans,
essentially repurposing it to function as a nectar sweetness receptor.[138] This adaptation for taste enabled
hummingbirds to detect and exploit sweet nectar as an energy source, facilitating their distribution across
geographical regions where nectar-bearing flowers are available.[138]

Tongue as a micropump

Hummingbirds drink with their long tongues by rapidly lapping nectar. Their tongues have tubes which run
down their lengths and help the hummingbirds drink the nectar.[140] While capillary action was believed to
be what drew nectar into these tubes, high-speed photography has revealed that the tubes open down their
sides as the tongue goes into the nectar, and then close around the nectar, trapping it so it can be pulled back
into the beak.[141][142] The tongue, which is forked, is compressed until it reaches nectar, then the tongue
springs open, the rapid action traps the nectar and the nectar moves up the grooves, like a pump action,
with capillary action not involved.[140][143] Consequently, tongue flexibility enables accessing, transporting
and unloading nectar.[144][145]

Feeders and artificial nectar

In the wild, hummingbirds visit flowers for food, extracting nectar,

which is 55% sucrose, 24% glucose, and 21% fructose on a dry-
matter basis.[146] Hummingbirds also take sugar-water from bird
feeders, which allow people to observe and enjoy hummingbirds
up close while providing the birds with a reliable source of energy,
especially when flower blossoms are less abundant. A negative
aspect of artificial feeders, however, is that the birds may seek less
flower nectar for food, and so may reduce the amount of pollination
their feeding naturally provides.[147]
Male ruby-throated hummingbird
White granulated sugar is used in hummingbird feeders in a 25% displaying his tongue
concentration as a common recipe,[148] although hummingbirds
will defend feeders more aggressively when sugar content is at
35%, indicating preference for nectar with higher sugar content.[149] Organic and "raw" sugars contain
iron, which can be harmful,[150] and brown sugar, agave syrup, molasses, and artificial sweeteners also
should not be used.[151] Honey is made by bees from the nectar of flowers, but it is not good to use in
feeders because when it is diluted with water, microorganisms easily grow in it, causing it to spoil
rapidly.[152][153][154]
Red food dye was once thought to be a favorable ingredient for the
nectar in home feeders, but it is unnecessary.[155] Commercial
products sold as "instant nectar" or "hummingbird food" may also
contain preservatives or artificial flavors, as well as dyes, which are
unnecessary and potentially harmful.[155][156] Although some
commercial products contain small amounts of nutritional additives,
hummingbirds obtain all necessary nutrients from the insects they
eat, rendering added nutrients unnecessary.[123]
Hummingbirds hovering at an
artificial nectar feeder

Visual cues of foraging

Hummingbirds have exceptional visual acuity providing them with discrimination of food sources while
foraging.[47] Although hummingbirds are thought to be attracted to color while seeking food, such as red
flowers or artificial feeders, experiments indicate that location and flower nectar quality are the most
important "beacons" for foraging.[157][158] Hummingbirds depend little on visual cues of flower color to
beacon to nectar-rich locations, but rather they used surrounding landmarks to find the nectar
reward.[159][160]

In at least one hummingbird species – the green-backed firecrown (Sephanoides sephaniodes) – flower
colors preferred are in the red-green wavelength for the bird's visual system, providing a higher contrast
than for other flower colors.[161] Further, the crown plumage of firecrown males is highly iridescent in the
red wavelength range (peak at 650 nanometers), possibly providing a competitive advantage of dominance
when foraging among other hummingbird species with less colorful plumage.[161] The ability to
discriminate colors of flowers and plumage is enabled by a visual system having four single cone cells and
a double cone screened by photoreceptor oil droplets which enhance color discrimination.[157][161]

Olfaction

While hummingbirds rely primarily on vision and hearing to assess competition from bird and insect
foragers near food sources, they may also be able to detect by smell the presence in nectar of insect
defensive chemicals (such as formic acid) and aggregation pheromones of foraging ants, which discourage
feeding.[162] Another study showed that hummingbirds are attracted to the nectar aroma of certain flowers
by the combination of volatile monoterpenes, including d-limonene, β-myrcene, and β-ocimene.[163]

Superficially similar species

Some species of sunbirds of Africa, southern and southeastern Asia, and Australia resemble hummingbirds
in appearance and behavior, as do perhaps also the honeyeaters of Australia and Pacific islands. These two
groups, however, are not related to hummingbirds, as their resemblance is due to convergent evolution.[164]

The hummingbird moth is often mistaken for a hummingbird.

In myth and culture

Aztecs wore hummingbird talismans, both artistic representations of hummingbirds and
fetishes made from actual hummingbird parts: emblematic for their vigor, energy, and
propensity to do work along with their sharp beaks that symbolically mimic instruments of
weaponry, bloodletting, penetration, and intimacy. Hummingbird talismans were prized as
drawing sexual potency, energy, vigor, and skill at arms and warfare to the wearer.[165]
 The Aztec god of war Huitzilopochtli is often depicted as
a hummingbird. It was also believed that fallen warriors
would return to earth as hummingbirds and
butterflies.[166] The Nahuatl word huitzil (hummingbird)
is an onomatopoeic word derived from the sounds of the
hummingbird's wing-beats and zooming flight.
One of the Nazca Lines depicts a hummingbird (right).
The Hopi and Zuni cultures have a hummingbird
Nazca Lines Hummingbird
creation myth about a young brother and sister who are
starving because drought and famine have come to the
land. Their parents have left to find food, so the boy
carves a piece of wood into a small bird to entertain his
sister. When the girl tosses the carving into the air, the
bird comes to life, turning into a hummingbird. The small
bird then flies to the God of Fertility and begs for rain,
and the god obliges the request, which helps the crops
to grow again.[167]
Trinidad and Tobago, known as "The land of the
hummingbird," displays a hummingbird on that nation's
Hummingbird theme on the
coat of arms,[168] 1-cent coin[169] and emblem of its
Caribbean Airlines Boeing 737
national airline, Caribbean Airlines (right).
The Gibson Hummingbird is an acoustic guitar
model/series produced by the Gibson Guitar Corporation.
During the National Costume competition of the Miss Universe 2016 beauty pageant, Miss
Ecuador Connie Jiménez wore a costume inspired by the hummingbirds of her land[170] that
included golden wings supposed to follow the movements of her arms.[171][172] However, it
accidentally got damaged during the dress rehearsal, and she appeared onstage with a
broken, drooping left wing.[173][174]

Gallery


Play media

Female ruby- Hummingbird Hummingbird with

throated Hummingbirds feeding from a yellow pollen on its
hummingbird feeding at 1500 fps flower in the beak in the
perched in Hudson, University of University of
Ohio California Botanical California Botanical
Garden Garden

Talamanca Costa's Juvenile Anna's Calypte anna

hummingbird hummingbird hummingbird with perched
tongue sticking out

Hummingbird chicks Hummingbird adult A female Anna's

in a nest in a cactus in its nest in a cactus hummingbird
in Mesa, Arizona in Mesa, Arizona perched on a small
branch

See also
AeroVironment Nano Hummingbird – artificial hummingbird
Macroglossum stellatarum – hummingbird hawk-moth
Hemaris – sphinx moths (hummingbird moths) confused with hummingbirds
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External links
The Hummingbird Website ([Link] Hummingbird photos, videos,
articles, links, frequently asked questions
High-resolution photo gallery of almost 100 species ([Link]
[Link]?q=gallery&g2_itemId=2182)
High-resolution photo gallery of many species of hummingbirds ([Link]
[Link]/[Link])
Video of hummingbird tongue acting as a micropump during nectar feeding ([Link]
[Link]/web/20170301202153/[Link]

Retrieved from "[Link]

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