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Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about species and genera , [1] they occur from Alaska to Tierra del Fuego , but the vast majority of the species are found in the tropics. They are small birds, with most species measuring 7. The smallest extant hummingbird species is the 5 cm 2.

The largest hummingbird species is the 23 cm 9. 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 because of the humming sound created by their beating wings , which flap at high frequencies audible to humans.

They hover in mid-air at rapid wing-flapping rates, which vary from around 12 beats per second in the largest species to around 80 per second in small hummingbirds.

Hummingbirds have the highest mass-specific metabolic rate of any homeothermic animal. The family Trochilidae was introduced in by Irish zoologist Nicholas Aylward Vigors with Trochilus as the type genus.

Molecular phylogenetic studies determined the relationships between the major groups of hummingbirds. Phaethornithinae — hermits. Patagoninae — giant hummingbird. 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. Around 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 hummingbird Patagona gigas , the mountaingems , the bees, and the emeralds. The hummingbird family has the third-greatest number of species of any bird family after the tyrant flycatchers and the tanagers.

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 , Gerald Mayr identified two 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.

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.

In , a million-year-old bird fossil unearthed in Wyoming was found to be a predecessor to both hummingbirds and swifts before the groups diverged. 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. A map of the hummingbird family tree — reconstructed from analysis of of the world's known species — shows rapid diversification from 22 million years ago.

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 hummingbird species can coexist in a specific region, such as the Andes range.

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

Within the same geographic region, hummingbird clades co-evolved with nectar-bearing plant clades, affecting mechanisms of pollination. Hummingbirds exhibit sexual size dimorphism according to Rensch's rule , [27] in which males are smaller than females in small-bodied species, and males are larger than females in large-bodied species. Sexual size and bill differences likely evolved due to constraints imposed by courtship, because mating displays of male hummingbirds require complex aerial maneuvers.

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.

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. Hummingbirds are specialized nectarivores [31] 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. Even in the most specialized hummingbird—plant mutualisms, though, the number of food plant lineages of the individual hummingbird species increases with time.

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. Hummingbirds and the plants they visit for nectar have a tight co-evolutionary association, generally called a plant—bird mutualistic network. These associations are also observed when closely related hummingbirds, for example two species of the same genus, visit distinct sets of flowering species.

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. Hummingbirds are named for the prominent humming sound their wingbeats make while flying and hovering to feed or interact with other hummingbirds. 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.

During turbulent airflow conditions created experimentally in a wind tunnel , hummingbirds exhibit stable head positions and orientation when they hover at a feeder. 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.

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.

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. Hummingbirds are highly sensitive to stimuli in their visual fields, responding to even minimal motion in any direction by reorienting themselves in midflight. 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.

Hummingbirds can use newly ingested sugars to fuel hovering flight within 30—45 minutes of consumption. A 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.

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.

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. 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.

The dynamic range of metabolic rates in hummingbirds [67] requires a parallel dynamic range in kidney function. 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.

Consisting of chirps, squeaks, whistles and buzzes, [72] hummingbird songs originate from at least seven specialized nuclei in the forebrain. The blue-throated hummingbird 's song differs from typical oscine songs in its wide frequency range, extending from 1. The avian vocal organ, the syrinx , plays an important role in understanding hummingbird song production.

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 torpor, to prevent dehydration , the kidney function declines, preserving needed compounds, such as glucose , water, and nutrients. 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. A hummingbird with a higher fat percentage will be less likely to enter a state of torpor compared to one with less fat, as a bird can use the energy from its fat stores.

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. Praying mantises have been observed as predators of hummingbirds.

As far as is known, male hummingbirds do not take part in nesting. Many hummingbird species use spider silk and lichen to bind the nest material together and secure the structure. Hummingbird building a nest in San Diego Zoo , video. Nest with two eggs in San Jose, California. Nest with two nestlings in Santa Monica, California.

Feeding two nestlings in Grand Teton National Park. Fallen Anna's hummingbird nest in Ventura County, California , shown next to a toothpick for scale. To serve courtship and territorial competition , many male hummingbirds have plumage with bright, varied coloration [98] 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.

By merely shifting position, feather regions of a muted-looking bird can instantly become fiery red or vivid green. 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. 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 wind tunnel used particle image velocimetry techniques to investigate the lift generated on the bird's upstroke and downstroke.

Many earlier studies had assumed that lift was generated equally during the two phases of the wingbeat cycle, as is the case of insects of a similar size.

Because of their flying technique, these birds no longer have an alula. As air density decreases, for example, at higher altitudes, the 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. 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.

For instance, it is about twice the diving speed of peregrine falcons in pursuit of prey. The outer tail feathers of male Anna's Calypte anna and Selasphorus hummingbirds e. 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 , [] 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.