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Hummingbirds are
birds Birds are a group of warm-blooded vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With ...

birds
native to the
Americas The Americas (also collectively called America) is a landmass comprising the totality of North America, North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...

Americas
and comprise the
biological family Family ( la, familia, plural ') is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy; it is classified between order (biology), order and genus. A family may be divided into subfamily, subfamilies, which are intermediate r ...
Trochilidae. With about 360 species, they occur from Alaska to
Tierra del Fuego #REDIRECT Tierra del Fuego #REDIRECT Tierra del Fuego#REDIRECT Tierra del Fuego Tierra del Fuego (, ; Spanish for "Land of Fire", formerly also Fireland in English) is an archipelago An archipelago ( ), sometimes called an island group or i ...

Tierra del Fuego
, but the vast majority of the species are found in the
tropics The tropics are the region of Earth Earth is the third planet from the Sun and the only astronomical object known to harbour and support life. 29.2% of Earth's surface is land consisting of continents and islands. The remaining 70.8% ...

tropics
. They are small birds, with most species measuring in length. The smallest
extant Extant is the opposite of the word extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the endling, last individual o ...
hummingbird species is the
bee hummingbird#REDIRECT Bee hummingbird {{Redirect category shell, 1= {{R from move {{R from other capitalisation ...
, which weighs less than . The largest hummingbird species is the
giant hummingbird The giant hummingbird (''Patagona gigas'') is the only member of the genus ''Patagona'' and the largest member of the hummingbird family, weighing and having a wingspan of approximately and length of . This is approximately the same length as a ...
, weighing . They are specialized for feeding on flower nectar, but all species also consume flying insects or spiders. Hummingbirds split from their
sister group In phylogenetics, a sister group or sister taxon comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and taxon B are sister groups to eac ...
, the swifts and
treeswift Treeswifts or crested swifts are a family (biology), family, Hemiprocnidae, of aerial near passerine birds, closely related to the Swift (bird), true swifts. The family contains a single genus, ''Hemiprocne'', with four species. They are distribute ...
s, 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 A wing is a type of fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust Thrust is a reaction (physics), reaction force (physics), force describ ...
, 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. Of those species that have been measured during flying in wind tunnels, their top speeds exceed . During
courtship Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...

courtship
, some male species dive from of height above a female at speeds around . Hummingbirds have the highest mass-specific metabolic rate of any
homeothermic Homeothermy, homothermy or homoiothermy is thermoregulation Thermoregulation is the ability of an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system ...
animal. To conserve energy when food is scarce and nightly when not foraging, they can go into
torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature Thermoregulation is the ability of an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμό ...
, a state similar to
hibernation Hibernation is a state of minimal activity and metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life Life is a characteristic that distinguishes physical entities that have biological ...

hibernation
, and slow their
metabolic rate Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main purposes of metabolism are: the conversion of the energy in food to energy available to run cell ...
to 1/15 of its normal rate.


Taxonomy and systematics

The family Trochilidae was introduced in 1825 by Irish zoologist
Nicholas Aylward Vigors Nicholas Aylward Vigors (1785 – 26 October 1840) was an Ireland, Irish zoologist and politician. He popularized the classification of birds on the basis of the quinarian system. Early life Vigors was born at Old Leighlin, County Carlow. He s ...
with ''
Trochilus The streamertails are hummingbirds in the genus ''Trochilus'', that are Endemism, endemic to Jamaica. It is the type genus of the family (biology), family Trochilidae. Today most authorities consider the two taxa in this genus as separate species ...
'' as the
type genus In biological taxonomy the type genus is the genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumsc ...
.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominately in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies of the hummingbirds have shown that the family is composed of nine major
clades A clade (; from grc, , ''klados'', "branch"), also known as a monophyletic group or natural group, is a group of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual cont ...
. When
Edward Dickinson Edward Dickinson (January 1, 1803 – June 16, 1874) was an American politician from Massachusetts Massachusetts (, ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England region of the Unite ...
and James Van Remsen, Jr. updated the ''
Howard and Moore Complete Checklist of the Birds of the World The ''Howard and Moore Complete Checklist of the Birds of the World'' is a book by Richard Howard and Alick Moore which presents a list of the bird species of the world. It was the first single-volume world bird list to include subspecies names, ...
'' for the 4th edition in 2013 they divided the hummingbirds into six
subfamilies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, ...
. The above
cladogram A cladogram (from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is appr ...

cladogram
of the hummingbird family is based on
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominately in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study by Jimmy McGuire and collaborators published in 2014. The English names are those introduced by Robert Bleiweiss, John Kirsch, and Juan Matheus in 1997. The Latin names are those proposed by Dickinson and Remsen in 2013. In traditional
taxonomy Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only about biological ...
, hummingbirds are placed in the order
Apodiformes Traditionally, the bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism ...
, which also contains the
swift The Society for Worldwide Interbank Financial Telecommunication (SWIFT), legally S.W.I.F.T. SCRL, is a Belgium, Belgian cooperative society that serves as an intermediary and executor of financial transactions between banks worldwide. It also ...
s, but some taxonomists have separated them into their own order, the Trochiliformes. Hummingbirds' wing
bone A bone is a rigid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphosa dubit ...

bone
s are hollow and fragile, making
fossil A fossil (from Classical Latin Classical Latin is the form of Latin language Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication used by humans, inc ...

fossil
ization 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 Russia ( rus, link=no, Россия, Rossiya, ), or the Russian Federation, is a country spanning Eastern Europe Eastern Europe is the eastern region of . There is no consistent definition of the precise area it covers, partly because th ...

Russia
today. Around 360 hummingbirds have been described. They have been traditionally divided into two
subfamilies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, ...
: the
hermits A hermit, or eremite (adjectival form In linguistics Linguistics is the science, scientific study of language. It encompasses the analysis of every aspect of language, as well as the methods for studying and modeling them. The trad ...
(subfamily Phaethornithinae) and the typical hummingbirds (subfamily
Trochilinae Trochilinae is one of the six subfamily, subfamilies that make up the hummingbird family (biology), family Trochilidae. The subfamily is divided into three Tribe (biology), tribes: Lampornithini (mountain gems) containing 18 species, Mellisugini ...
, all the others). Molecular phylogenetic studies have shown, though, that the hermits are
sister A sister is a woman A woman is an adult female Female (symbol: ♀) is the sex of an organism, or a part of an organism, that produces non-mobile ovum, ova (egg cells). Barring rare medical conditions, most female mammals, including fema ...
to the topazes, making the former definition of the Trochilinae not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

monophyletic
. The hummingbirds form nine major
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

clade
s: the topazes and
jacobins , logo = JacobinVignette03.jpg , logo_size = 180px , logo_caption = Seal of the Jacobin Club (1792–1794) , motto = "Live free or die"(french: Vivre libre ou mourir) , successor = Pa ...
, the hermits, the
mangoes A mango is an edible stone fruit ), showing both fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which angiosper ...
, the coquettes, the brilliants, the
giant hummingbird The giant hummingbird (''Patagona gigas'') is the only member of the genus ''Patagona'' and the largest member of the hummingbird family, weighing and having a wingspan of approximately and length of . This is approximately the same length as a ...
(''Patagona gigas''), the mountaingems, the bees, and the emeralds. The topazes and jacobins combined have the oldest split with the rest of the hummingbirds. The hummingbird family has the third-greatest number of species of any bird family (after the
tyrant flycatcher The tyrant flycatchers (Tyrannidae) are a Family (biology), family of passerine birds which occur throughout North America, North and South America. They are considered the largest family of birds known to exist in the world, with more than 400 ...
s and the
tanager The tanagers (singular ) comprise the bird family (biology), family Thraupidae, in the order Passeriformes. The family has a Neotropical distribution and is the second-largest family of birds. It represent about 4% of all avian species and 12% ...
s).
Fossil A fossil (from Classical Latin Classical Latin is the form of Latin language Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication used by humans, inc ...

Fossil
hummingbirds are known from the
Pleistocene The Pleistocene ( , often referred to as the ''Ice Age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the earth’s most recent period of repeated glaciations. Before a change finally ...
of
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At 8.5 million square kilometers (3.2 million square miles) and with over 211 mill ...

Brazil
and the
Bahamas The Bahamas (), known officially as The Commonwealth of The Bahamas, is a sovereign country within the Lucayan Archipelago of the West Indies in the Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the a ...

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 Gerald Mayr is a German paleontology, palaeontologist who is Curator of Ornithology at the Naturmuseum Senckenberg, Senckenberg Research Institute in Frankfurt, Frankfurt am Main, Hesse. He has published extensively on fossil birds, especially the ...
identified two 30-million-year-old hummingbird fossils. The fossils of this primitive hummingbird species, named ''
Eurotrochilus ''Eurotrochilus'' is a genus of extinct members of the stem group Trochilidae and are the closest relatives of the crown group Trochilidae, which includes modern hummingbirds. Despite ''Eurotrochilus'' having looked very similar to modern hummingb ...
inexpectatus'' ("unexpected European hummingbird"), had been sitting in a
museum A museum ( ; plural museums or, rarely, musea) is a building or institution that cares for and displays a collection Collection or Collections may refer to: * Cash collection, the function of an accounts receivable department * Collec ...

museum
drawer in
Stuttgart Stuttgart (; Swabian: ; ) is the capital Capital most commonly refers to: * Capital letter Letter case (or just case) is the distinction between the letters that are in larger uppercase or capitals (or more formally ''majuscule'') ...

Stuttgart
; they had been unearthed in a clay pit at
Wiesloch Wiesloch (, locally ), is a town in northern Baden-Württemberg Baden-Württemberg (; ) is a state (''Land'') in southwest Germany Southern Germany () as a region has no exact boundary but is generally taken to include the areas in which Up ...

Wiesloch
–Frauenweiler, south of
Heidelberg Heidelberg () is a university town in the German state The Federal Republic of Germany, as a federal state, consists of sixteen partly sovereign federated states (german: Land (state), plural (states); commonly informally / federated s ...

Heidelberg
,
Germany ) , image_map = , map_caption = , map_width = 250px , capital = Berlin Berlin (; ) is the Capital city, capital and List of cities in Germany by population, largest city of Germany by both area and population. Its 3,769,495 inh ...

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 The Messel Pit (german: Grube Messel) is a disused quarry Image:0 Carrières du Hainaut à Soignies (2).JPG, Stone quarry in Soignies, Hainaut (province), Belgium A quarry is a type of open-pit mining, open-pit mine in which dimension s ...
and in the
Caucasus The Caucasus (), or Caucasia (), is a region spanning Europe and Asia. It is situated between the Black Sea and the Caspian Sea and mainly occupied by Armenia, Azerbaijan, Georgia (country), Georgia, and parts of Southern Russia. It is home to ...
, 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 Climate is the long-term pattern of weather Weather is the state of the atmosphere An atmosphere (from the greek words ἀτμός ''(atmos)'', meaning 'vapour', and σφαῖρα ''(sphaira)'', meaning 'ball' or 'sphere') is a la ...

climate
quite similar to that of the northern
Caribbean The Caribbean (, ; es, Caribe; french: Caraïbes; ht, Karayib; also gcf, label=Antillean Creole Antillean Creole (Antillean French Creole, Kreyol, Kwéyòl, Patois) is a French-based creole, which is primarily spoken in the Lesser Antilles ...
or southernmost
China China (), officially the People's Republic of China (PRC; ), is a country in East Asia East Asia is the eastern region of Asia Asia () is Earth's largest and most populous continent, located primarily in the Eastern Hemisphere ...

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 ''Eurotrochilus'' is a genus of extinct members of the stem group Trochilidae and are the closest relatives of the crown group Trochilidae, which includes modern hummingbirds. Despite ''Eurotrochilus'' having looked very similar to modern hummingb ...
'' and the modern fossils. The astounding morphological
adaptation In , adaptation has three related meanings. Firstly, it is the dynamic evolutionary process that fits s to their environment, enhancing their . Secondly, it is a state reached by the population during that process. Thirdly, it is a or adapti ...

adaptation
s, the decrease in size, and the dispersal to the Americas and extinction in
Eurasia Eurasia () is the largest continent A continent is any of several large landmass A landmass, or land mass, is a large region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a ...

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 The Miocene ( ) is the first geological epoch In geochronology, an epoch is a subdivision of the geologic timescale that is longer than an age (geology), age but shorter than a period (geology), period. The current epoch is the Holocene Epoch of ...
, some 12 to 13 million years ago, during the uplifting of the northern
Andes The Andes, Andes Mountains or Andean Mountains ( es, Cordillera de los Andes) are the List of mountain ranges#Mountain ranges by length, longest continental mountain range in the world, forming a continuous highland along the western edge of Sou ...

Andes
. In 2013, a 50-million-year-old bird fossil unearthed in
Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. The List of U.S. states and territories by area, 10th largest state by area, it is also the List of U.S. states and territories b ...
was found to be a predecessor to both hummingbirds and swifts before the groups diverged.


Lists of genera and species

* List of hummingbird species, sortable alphabetically by common name, binomial name, or taxonomic sequence


Evolution

Hummingbirds are thought to have split from other members of Apodiformes, the insectivorous swifts (family Apodidae) and
treeswift Treeswifts or crested swifts are a family (biology), family, Hemiprocnidae, of aerial near passerine birds, closely related to the Swift (bird), true swifts. The family contains a single genus, ''Hemiprocne'', with four species. They are distribute ...
s (family Hemiprocnidae) about 42 million years ago, probably in Eurasia. Despite their current New World distribution, the earliest known species of humming bird are known from the early
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define ...
(
Rupelian The Rupelian is, in the geologic timescale The geologic time scale (GTS) is a system of chronological datingChronological dating, or simply dating, is the process of attributing to an object or event a date in the past, allowing such object or ...
~34-28 million years ago) of Europe, belonging to the genus ''
Eurotrochilus ''Eurotrochilus'' is a genus of extinct members of the stem group Trochilidae and are the closest relatives of the crown group Trochilidae, which includes modern hummingbirds. Despite ''Eurotrochilus'' having looked very similar to modern hummingb ...
,'' which is very similar in its morphology to modern hummingbirds. 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. A map of the hummingbird family treereconstructed from analysis of 284 of the world's 338 known
species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individu ...

species
shows rapid diversification from 22 million years ago. Hummingbirds fall into nine main
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

clade
s, the topazes,
hermits A hermit, or eremite (adjectival form In linguistics Linguistics is the science, scientific study of language. It encompasses the analysis of every aspect of language, as well as the methods for studying and modeling them. The trad ...
,
mangoes A mango is an edible stone fruit ), showing both fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which angiosper ...
, brilliants, coquettes, the
giant hummingbird The giant hummingbird (''Patagona gigas'') is the only member of the genus ''Patagona'' and the largest member of the hummingbird family, weighing and having a wingspan of approximately and length of . This is approximately the same length as a ...
, mountaingems,
bees Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
, and
emeralds Emerald is a gemstone A gemstone (also called a gem, fine gem, jewel, precious stone, or semi-precious stone) is a piece of mineral crystal which, in cut and polished form, is used to make jewellery, jewelry or other adornments. However, cer ...
, defining their relationship to
nectar Nectar is a sugar Sugar is the generic name for , soluble s, many of which are used in food. Simple sugars, also called s, include , , and . Compound sugars, also called s or double sugars, are molecules made of two monosaccharides jo ...

nectar
-bearing
flowering plant Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greec ...

flowering plant
s and the birds' continued spread into new geographic areas. 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
evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, ...

evolution
ary pattern, as many as 140 hummingbird species can coexist in a specific region, such as the
Andes The Andes, Andes Mountains or Andean Mountains ( es, Cordillera de los Andes) are the List of mountain ranges#Mountain ranges by length, longest continental mountain range in the world, forming a continuous highland along the western edge of Sou ...

Andes
range Range may refer to: Geography * Range (geographic)A range, in geography, is a chain of hill A hill is a landform A landform is a natural or artificial feature of the solid surface of the Earth or other planetary body. Landforms together ...

range
. The hummingbird
evolutionary tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram A diagram is a symbolic representation Representation may refer to: Law a ...

evolutionary tree
shows one key evolutionary factor appears to have been an altered
taste receptor A taste receptor is a type of cellular receptor In biochemistry and pharmacology, receptors are chemical structures, composed of protein, that receive and Signal_transduction, transduce signals that may be integrated into biological systems. T ...
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. Hummingbirds remain in dynamic diversification inhabiting ecological regions across South America, North America, and the Caribbean, indicating an enlarging
evolutionary radiation An evolutionary radiation is an increase in taxonomic Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific class ...
. Within the same geographic region, hummingbird clades co-evolved with nectar-bearing plant clades, affecting mechanisms of
pollination Pollination is the transfer of pollen Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes (sperm cells). Pollen grains have a hard coat ...

pollination
. 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'').


Sexual dimorphisms

Hummingbirds exhibit sexual size dimorphism according to Rensch's rule, in which males are smaller than females in small-bodied species, and males are larger than females in large-bodied species. The extent of this sexual size difference varies among clades of hummingbirds. For example, the Mellisugini clade (bees) exhibits a large size dimorphism, with females being larger than males. Conversely, the Lesbiini clade (coquettes) displays very little size dimorphism; males and females are similar in size. Sexual dimorphisms in
bill Bill(s) may refer to: Common meanings * Banknote A banknote (often known as a bill (in the US and Canada), paper money, or simply a note) is a type of negotiable instrument, negotiable promissory note, made by a bank or other licensed author ...

bill
size and shape are also present between male and female hummingbirds, where in many clades, females have longer, more curved bills favored for accessing nectar from tall flowers. For males and females of the same size, females tend to have larger bills. Sexual size and bill differences likely evolved due to constraints imposed by courtship, because mating displays of male hummingbirds require complex aerial maneuvers. Males tend to be smaller than females, allowing conservation of energy to
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing In agriculture Agriculture is the science, art and practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentism, se ...

forage
competitively and participate more frequently in
courtship Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...
. Thus,
sexual selection Sexual selection is a mode of natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrat ...
favors smaller male hummingbirds. 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. Thus, females are better at foraging, acquiring flower nectar, and supporting the energy demands of their larger body size.
Directional selection upThree types of selection In population genetics, directional selection, or positive selection is a mode of natural selection Natural selection is the differential survival and reproduction of individuals due to differences in pheno ...

Directional selection
thus favors the larger hummingbirds in terms of acquiring food. 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. Depending on which sex holds territory in the species, the other sex having a longer bill and being able to feed on a wide variety of flowers is advantageous, decreasing
intraspecific competition Intraspecific competition is an interaction in population ecology, whereby members of the same species compete for limited resources. This leads to a reduction in Fitness (biology), fitness for both individuals, but the more fit individual survives ...
. For example, in species of hummingbirds where males have longer bills, males do not hold a specific territory and have a
lek mating A lek is an aggregation of male animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic materia ...
system. 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.


Co-evolution with ornithophilous flowers

Hummingbirds are specialized
nectarivore In zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, ...
s 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 Corolla may refer to: *Corolla (botany) upright=1.4, Diagram showing the parts of a mature flower. In this example the perianth is separated into a calyx (sepals) and corolla (petals) The perianth (perigonium, perigon or perigone in monocots) is ...

corolla
length, curvature, and volume. 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. Even in the most specialized hummingbird-plant mutualisms, though, the number of food plant lineages of the individual hummingbird species increases with time. The bee hummingbird (''Mellisuga helenae'') – the world's smallest bird – evolved to
dwarfism Dwarfism occurs when an organism is exceptionally small. In humans, it is sometimes defined as Short stature, an adult height of less than , regardless of sex; the average adult height among people with dwarfism is , although some individuals ...
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. Many plants
pollinated Pollination is the transfer of pollen from a male part of a plant to a female part of a plant, later enabling fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and im ...

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
wavelength In physics Physics is the that studies , its , its and behavior through , and the related entities of and . "Physical science is that department of knowledge which relates to the order of nature, or, in other words, to the regular su ...

wavelength
s into the near-ultraviolet, but hummingbird-pollinated flowers do not reflect these wavelengths as many insect-pollinated flowers do. This narrow
color spectrum Laser beams with visible spectrum The visible spectrum is the portion of the electromagnetic spectrum The electromagnetic spectrum is the range of frequencies (the spectrum A spectrum (plural ''spectra'' or ''spectrums'') is a condition ...
may render hummingbird-pollinated flowers relatively inconspicuous to most insects, thereby reducing
nectar robbing Nectar robbing is a foraging behavior utilized by some organisms that feed on floral nectar. "Nectar robbers" usually feed from holes bitten in flowers, rather than by entering through the flowers' natural openings. Often, nectar robbers avoid con ...

nectar robbing
. Hummingbird-pollinated flowers also produce relatively weak nectar (averaging 25% sugars w/w) containing a high proportion of
sucrose Sucrose is a type of sugar Sugar is the generic name for , soluble s, many of which are used in food. Simple sugars, also called s, include , , and . Compound sugars, also called s or double sugars, are molecules made of two monosacchari ...

sucrose
, whereas insect-pollinated flowers typically produce more concentrated nectars dominated by
fructose Fructose, or fruit sugar, is a ketonic simple sugar Monosaccharides (from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a cou ...

fructose
and
glucose Glucose is a simple with the . Glucose is the most abundant , a subcategory of s. Glucose is mainly made by and most during from water and carbon dioxide, using energy from sunlight, where it is used to make in s, the most abundant carbohydr ...

glucose
. Hummingbirds and the plants they visit for nectar have a tight co-evolutionary association, generally called a plant–bird mutualistic network. 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.


Bill evolution

Upon maturity, males of a particular species, ''Phaethornis longirostris,'' the long-billed hermit, appear to be evolving a
dagger A dagger is a knife A knife (plural knives; from Old Norse 'knife, dirk') is a tool or weapon with a cutting edge or blade, often attached to a handle or hilt. One of the earliest tools used by humanity, knives appeared at least Stone Ag ...

dagger
-like weapon on the
beak The beak, bill, and/or rostrum is an external anatomical structure found mostly in birds Birds are a group of warm-blooded vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellu ...

beak
tip as a secondary sexual trait to defend mating areas.


Specialized characteristics and metabolism


Humming

A hummingbird is named for the prominent humming sound its wingbeats make while flying and hovering to feed or interact with other hummingbirds. Humming serves communication purposes by alerting other birds of the arrival of a fellow forager or potential mate. The humming sound derives from
aerodynamic force An aerodynamic force is a force exerted on a body by the air (or other gas) in which the body is immersed, and is due to the relative motion The ''relative velocity'' \vec_ (also \vec_ or \vec_) is the velocity of an object or observer B in the r ...
s generated by both the downstrokes and upstrokes of the rapid wingbeats, causing that evoke an acoustic quality likened to that of a
musical instrument A musical instrument is a device created or adapted to make musical sounds. In principle, any object that produces sound can be considered a musical instrument—it is through purpose that the object becomes a musical instrument. A person who play ...
. The humming sound of hummingbirds is unique among flying animals, compared to the whine of
mosquito Mosquitoes are members of a group of almost 3,600 species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defin ...

mosquito
es, buzz of
bee Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...

bee
s, and 'whoosh' of larger birds. 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 pectoralis major () is a thick, fan-shaped or triangular convergent muscle, situated at the chest of the human body. It makes up the bulk of the chest muscles Skeletal muscles (commonly referred to as muscles) are Organ (biology), organs ...

pectoralis major
(the main downstroke muscle) and supracoracoideus (the main upstroke muscle).


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. 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. During turbulent airflow conditions created experimentally in a
wind tunnel Wind tunnels are large tubes with air blowing through them which are used to replicate the interaction between air and an object flying through the air or moving along the ground. Researchers use wind tunnels to learn more about how an aircraft ...

wind tunnel
, hummingbirds exhibit stable head positions and orientation when they hover at a feeder. When wind gusts from the side, hummingbirds compensate by increasing wing-stroke
amplitude The amplitude of a period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Period, a descriptor for ...

amplitude
and stroke plane angle and by varying these parameters asymmetrically between the wings and from one stroke to the next. They also vary the orientation and enlarge the collective
surface area The surface area of a solid Solid is one of the four fundamental states of matter 4 (four) is a number A number is a mathematical object A mathematical object is an abstract concept arising in mathematics. In the usual language of ...

surface area
of their tail feathers into the shape of a
fan Fan commonly refers to: * Fan (machine), a machine for producing airflow, often for cooling ** Hand fan, an implement held and waved by hand to move air * Fan (person), short for fanatic; an enthusiast or supporter, especially with regard to enterta ...

fan
. While hovering, the
visual system The visual system comprises the sensory organ A sense is a biological system A biological system is a complex network which connects several biologically relevant entities. Biological organization spans several scales and are determined b ...
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. 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.


Vision

Although hummingbird eyes are small in diameter (5–6 mm), they are accommodated in the skull by reduced skull
ossification Ossification (or osteogenesis) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in the formation of no ...
, and occupy a relatively larger proportion of the skull compared to other birds and animals. Further, hummingbird eyes have relatively large
cornea The cornea is the transparent Transparency, transparence or transparent most often refer to transparency and translucency, the physical property of allowing the transmission of light through a material. They may also refer to: Literal uses * ...

cornea
s, which comprise about 50% of the total transverse eye diameter, combined with an extraordinary density of
retinal ganglion cell A retinal ganglion cell (RGC) is a type of neuron located near the inner surface (the ganglion cell layer) of the retina of the human eye, eye. It receives visual information from photoreceptor cell, photoreceptors via two intermediate neuron typ ...
s responsible for visual processing, containing some 45,000
neuron A neuron or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapse In the nervous system In biology Biology is the natural science that studies life and living ...

neuron
s per mm2. The enlarged cornea relative to total eye diameter serves to increase the amount of light perception by the eye when the
pupil The pupil is a black hole located in the center of the iris of the eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo ...

pupil
is dilated maximally, enabling
nocturnal Nocturnality is an animal behavior Ethology is the scientific Science (from the Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was original ...
flight. 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 physics Physics is the natural science that studies matter, its Elementary particle, fundamental constituents, its Motion (physics), motion and behavior through Spacetime, space and time, and the related entities of energy and force. "Phy ...
in the lateral and frontal
visual field The visual field is the "spatial array of visual sensations available to observation in introspection Introspection is the examination of one's own conscious thoughts and feelings. In psychology, the process of introspection relies on the observ ...
s. Morphological studies of the hummingbird brain showed that
neuron A neuron or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapse In the nervous system In biology Biology is the natural science that studies life and living ...

neuron
al
hypertrophy __NOTOC__ Hypertrophy (, from Greek ὑπέρ "excess" + τροφή "nourishment") is the increase in the volume of an organ or tissue due to the enlargement of its component Cell (biology), cells. It is distinguished from hyperplasia, in which ...

hypertrophy
relatively the largest in any bird exists in a region called the pretectal nucleus lentiformis (called the ''nucleus of the
optic tract The optic tract (from the Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to ...
'' in mammals) responsible for refining dynamic visual processing while hovering and during rapid flight. 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 Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...

courtship
. A study of
broad-tailed hummingbird The broad-tailed hummingbird (''Selasphorus platycercus'') is a medium-sized hummingbird Hummingbirds are birds native to the Americas and constitute the biological family Trochilidae. There are about 360 species. They occur from Alaska to ...
s indicated that hummingbirds have a fourth color-sensitive visual cone (humans have three) that detects
ultraviolet light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength In physics Physics (from grc, φυσική (ἐπιστήμη), physikḗ (epistḗmē), knowledge of nature, from ''phýsis'' 'nature'), , is the natural s ...

ultraviolet light
and enables discrimination of non-spectral colors, possibly having a role in
courtship display A courtship display is a set of display behaviors Image:Feather of male Pavo cristatus (Indian peafowl).jpg, 250px, Many male birds have brightly coloured plumage for display. This feather is from a male Indian peafowl ''Pavo cristatus''. Displa ...
s, territorial defense, and predator evasion. The fourth color cone would extend the range of visible colors for hummingbirds to perceive ultraviolet light and color combinations of feathers and
gorget A gorget , from the French ' meaning throat In vertebrate anatomy, the throat is the front part of the neck, positioned in front of the vertebra. It contains the Human pharynx, pharynx and larynx. An important section of it is the epiglotti ...

gorget
s, 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. Hummingbirds are highly sensitive to stimuli in their visual fields, responding to even minimal motion in any direction by reorienting themselves in midflight. Their visual sensitivity allows them to precisely hover in place while in complex and dynamic natural environments, functions enabled by the
lentiform nucleus The lentiform nucleus, or lenticular nucleus, comprises the putamen The putamen (; from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoke ...

lentiform nucleus
which is tuned to fast-pattern velocities, enabling highly tuned control and collision avoidance during forward flight.


Metabolism

With the exception of insects, hummingbirds while in flight have the highest
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life Life is a characteristic that distinguishes physical entities A bubble of exhaled gas in water In common usage and classical mechanics, a phys ...

metabolism
of all animals – a necessity to support the rapid beating of their wings during hovering and fast forward flight. Their heart rate can reach as high as 1,260 bpm, a rate once measured in a blue-throated hummingbird, with a
breathing rate The respiratory rate is the rate at which breathing occurs. This is usually measured in breaths per minute and is set and controlled by the respiratory centre. Measurement The respiratory rate in humans is measured by counting the number of brea ...
of 250 bpm, even at rest. During flight, oxygen consumption per gram of muscle tissue in a hummingbird is about 10 times higher than that measured in elite human athletes. Hummingbirds are rare among vertebrates in their ability to rapidly make use of ingested sugars to fuel energetically expensive hovering flight, 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. 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 ATP may refer to: Companies and organizations * Association of Tennis Professionals * American Technical Publishers * ', a Danish pension * Armenia Tree Project * Association for Transpersonal Psychology * ATP architects engineers office * ATP ...

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 Fasting is the willful refrainment from eating Eating (also known as consuming) is the ingestion Ingestion is the consumption of a substance by an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ...

fasting
or to power migratory flights. Studies of hummingbird metabolism address how a migrating
ruby-throated hummingbird The ruby-throated hummingbird (''Archilochus colubris'') is a species of hummingbird Hummingbirds are birds native to the Americas and constitute the biological family Trochilidae. There are about 360 species. They occur from Alaska to Tier ...

ruby-throated hummingbird
can cross of the
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin 400px, Diagrammatic cross-section of an ocean basin, showing the various geographic features In hydrology Hydrology (from Greek: wikt:ὕδωρ, ὕδωρ, "hýdōr" meaning ...

Gulf of Mexico
on a nonstop flight. 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.


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 Thermoregulation is the ability of an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxo ...
for heat dissipation, which becomes an even greater challenge in hot, humid climates. Hummingbirds dissipate heat partially by
evaporation Evaporation is a type of vaporization Vaporization (or vaporisation) of an element or compound is a phase transition from the liquid phase to vapor. There are two types of vaporization: evaporation and boiling. Evaporation is a surface phe ...

evaporation
through exhaled air, and from body structures with thin or no feather covering, such as around the eyes, shoulders, under the wings (
patagia The patagium (plural: patagia) is a membranous structure that assists an animal in gliding or flight Flight or flying is the process by which an object (physics), object motion (physics), moves through a space without contacting any planetary s ...
), and feet. While hovering, hummingbirds do not benefit from the heat loss by during forward flight, except for air movement generated by their rapid wing-beat, possibly aiding convective heat loss from the extended feet. Smaller hummingbird species, such as the
calliope In Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the Cosmogony, origin and Cosmology#Metaphysical cosmology, nature of ...
, appear to adapt their relatively higher surface-to-volume ratio to improve convective cooling from air movement by the wings. When air temperatures rise above , 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 Exhalation (or expiration) is the flow of the breath Breathing (or ventilation) is the process of moving air File:Atmosphere gas proportions.svg, Composition of Earth's atmosphere by volume, excluding water vapor. Lower pie represent ...
. In cold climates, hummingbirds retract their feet into breast feathers to eliminate skin exposure and minimize heat dissipation.


Kidney function

The dynamic range of metabolic rates in hummingbirds requires a parallel dynamic range in
kidney The kidneys are two reddish-brown bean-shaped organs An organ is a group of tissues with similar functions. Plant life and animal life rely on many organs that co-exist in organ systems. A given organ's tissues can be broadly categorized ...

kidney
function. 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. During brief periods of water deprivation, however, such as in nighttime torpor, GFR drops to zero, preserving body water. Hummingbird kidneys also have a unique ability to control the levels of
electrolyte An electrolyte is a medium containing ions that is electrically conducting Electrical resistivity (also called specific electrical resistance or volume resistivity) is a fundamental property of a material that quantifies how strongly it resis ...

electrolyte
s after consuming nectars with high amounts of
sodium Sodium is a chemical element In chemistry, an element is a pure Chemical substance, substance consisting only of atoms that all have the same numbers of protons in their atomic nucleus, nuclei. Unlike chemical compounds, chemical eleme ...

sodium
and
chloride The chloride ion An ion () is an atom An atom is the smallest unit of ordinary matter In classical physics and general chemistry, matter is any substance that has mass and takes up space by having volume. All everyday objects th ...

chloride
or none, indicating that kidney and glomerular structures must be highly specialized for variations in nectar
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
quality. Morphological studies on
Anna's hummingbird Anna's hummingbird (''Calypte anna'') is a medium-sized bird species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecu ...

Anna's hummingbird
kidneys showed adaptations of high
capillary A capillary is a small blood vessel The blood vessels are the components of the circulatory system The circulatory system, also called the cardiovascular system or the vascular system, is an organ system that permits blood to circulate ...

capillary
density in close proximity to
nephron The nephron is the minute or microscopic structural and functional unit of the kidney The kidneys are two reddish-brown bean-shaped organs An organ is a group of tissues with similar functions. Plant life and animal life rely on many or ...

nephron
s, allowing for precise regulation of water and electrolytes.


Song and vocal learning

Consisting of chirps, squeaks, whistles and buzzes, hummingbird songs originate from at least seven specialized
nuclei ''Nucleus'' (plural nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA ...
in the
forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hindbrain (rhombencephalon) are the three Brain ...
. A
genetic expression Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. These ...
study showed that these nuclei enable
vocal learningVocal learning is the ability to modify acoustic and syntactic sounds, acquire new sounds via imitation, and produce vocalizations. "Vocalizations" in this case refers only to sounds generated by the vocal organ (mammalian larynx or avian syrinx) as ...
(ability to acquire vocalizations through imitation), a rare trait known to occur in only two other groups of birds (
parrot Parrots, also known as psittacines , are birds of the roughly 398 species in 92 genus (biology), genera comprising the order (biology), order Psittaciformes , found mostly in tropics, tropical and subtropics, subtropical regions. The order is s ...

parrot
s and
songbird A songbird is a bird Birds are a group of warm-blooded vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biologi ...

songbird
s) and a few groups of mammals (including humans, , and
bat Bats are mammal Mammals (from Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the po ...

bat
s). Within the past 66 million years, only hummingbirds, parrots, and songbirds out of 23 bird
orders Orders is a surname In some cultures, a surname, family name, or last name is the portion of one's personal name 300px, First/given, middle and last/family/surname with John Fitzgerald Kennedy as example. This shows a structure typical f ...
may have independently evolved seven similar forebrain structures for singing and vocal learning, indicating that evolution of these structures is under strong
epigenetic In biology, epigenetics is the study of heritability, heritable phenotype changes that do not involve alterations in the DNA sequence. The Ancient Greek, Greek prefix ''wikt:epi-, epi-'' ( "over, outside of, around") in ''epigenetics'' implies f ...
constraints possibly derived from a common ancestor. 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. It also produces vocalizations which do not function in communication. 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. The avian vocal organ, the
syrinx In classical Greek mythology Greek mythology is the body of myth Myth is a folklore genre Folklore is the expressive body of culture shared by a particular group of people; it encompasses the tradition A tradition is a belief ...
, plays an important role in understanding hummingbird song production. What makes the hummingbird's syrinx different from that of other birds in the
Apodiformes Traditionally, the bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism ...
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.


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 Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature Thermoregulation is the ability of an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμό ...
) to prevent energy reserves from falling to a critical level. During nighttime torpor,
body temperature Thermoregulation is the ability of an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies the Life#Biology, properties of life. It is a sy ...
falls from 40 to 18 °C, with
heart The heart is a muscular MUSCULAR (DS-200B), located in the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain,Usage is mixed. The Guardian' and Telegraph' use ...

heart
and
breathing rate The respiratory rate is the rate at which breathing occurs. This is usually measured in breaths per minute and is set and controlled by the respiratory centre. Measurement The respiratory rate in humans is measured by counting the number of brea ...
s both slowed dramatically (heart rate of roughly 50 to 180 bpm from its daytime rate of higher than 1000 bpm). During torpor, to prevent
dehydration In physiology, dehydration is a lack of total body water In physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology Biology is the natural science that studie ...

dehydration
, the GFR ceases, preserving needed compounds such as glucose, water, and nutrients. 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. The circulating
hormone A hormone (from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is appr ...

hormone
,
corticosterone Corticosterone, also known as 17-deoxycortisol and 11β,21-dihydroxyprogesterone, is a 21-carbon steroid hormone A steroid hormone is a steroid that acts as a hormone. Steroid hormones can be grouped into two classes: corticosteroids (typically m ...

corticosterone
, is one signal that arouses a hummingbird from torpor. 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. The hummingbirds of the Andes in South America are known for entering exceptionally deep torpor and dropping their body temperature.


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 Fledging is the stage in a flying animal A number of animals are capable of aerial locomotion, either by powered flight or by gliding (flight), gliding. This trait has appeared by evolution many times, without any single ancestor. Flight has ...
, those that survive may occasionally live a decade or more. Among the better-known North American species, the typical lifespan is probably 3 to 5 years. For comparison, the smaller
shrew Shrews (family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or social territory, typically subject to ...
s, among the smallest of all mammals, seldom live longer than 2 years. 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. Other longevity records for banded hummingbirds include an estimated minimum age of 10 years 1 month for a female
black-chinned hummingbird The black-chinned hummingbird (''Archilochus alexandri'') is a small hummingbird occupying a broad range of habitats. It is migratory, spending winter as far south as Mexico Mexico ( es, México ; Nahuan languages: ), officially the United ...
similar in size to the broad-tailed hummingbird, and at least 11 years 2 months for a much larger .Patuxent Wildlife Research Center, Bird Banding Laboratory
Longevity Records AOU Numbers 3930–4920
2009-08-31. Retrieved 2009-09-27.


Predators

have been observed as predators of hummingbirds. Other predators include
dragonflies A dragonfly is a flying insect belonging to the order Odonata, infraorder Anisoptera (from Ancient Greek, Greek ἄνισος ''anisos'', "unequal" and πτερόν ''pteron'', "wing", because the hindwing is broader than the forewing). Adult d ...

dragonflies
,
frog A frog is any member of a diverse and largely carnivorous A carnivore , meaning "meat Meat is animal flesh that is eaten as food. Humans have hunted and killed animals for meat since prehistoric times. The advent of civilization all ...

frog
s,
orb-weaver spider Orb-weaver spiders are members of the spider family (biology), family Araneidae. They are the most common group of builders of spiral wheel-shaped spider web, webs often found in gardens, fields, and forests. The English word ''orb'' can mean "cir ...
s, and other birds, such as the
roadrunner The roadrunners (genus ''Geococcyx''), also known as chaparral birds or chaparral cocks, are two species of fast-running ground cuckoo The cuckoos are a family of bird Birds are a group of warm-blooded vertebrates constituting the cl ...

roadrunner
.


Reproduction

As far as is known, male hummingbirds do not take part in nesting. Most species build a cup-shaped nest on the branch of a tree or shrub, 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 A walnut is the Nut (fruit), nut of any tree of the genus ''Juglans'' (family Juglandaceae), particularly the Persian or English walnut, ''Juglans regia''. A walnut is the edible seed of a drupe, and thus not a true botanical nut. It is com ...

walnut
shell to several centimeters in diameter. Many hummingbird species use
spider silk Spider silk is a protein Proteins are large biomolecule , showing alpha helices, represented by ribbons. This poten was the first to have its suckture solved by X-ray crystallography by Max Perutz and Sir John Cowdery Kendrew in 1 ...

spider silk
and
lichen A lichen ( , ) is a composite organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecu ...

lichen
to bind the nest material together and secure the structure. The unique properties of the silk allow the nest to expand as the young hummingbirds grow.Archived a
Ghostarchive
and th
Wayback Machine
Two white eggs are laid, which despite being the smallest of all bird eggs are large relative to the adult hummingbird's size. lasts 14 to 23 days, depending on the species, ambient temperature, and female attentiveness to the nest. The mother feeds her nestlings on small
arthropod An arthropod (, (gen. ποδός)) is an invertebrate animal having an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Euarthropoda,Reference showing that Euarthropoda is a phylum: ...
s and nectar by inserting her bill into the open mouth of a
nestling Birds are a group of warm-blooded vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With ...
, and then regurgitating the food into its crop. 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. Trochilidae - Hummingbird.webm, Hummingbird building a nest in
San Diego Zoo The San Diego Zoo is a zoo in Balboa Park (San Diego), Balboa Park, San Diego, California, housing over 12,000 animals of more than 650 species and subspecies on of Balboa Park leased from the City of San Diego. Its parent organization, San Dieg ...

San Diego Zoo
, video Hummingbird Incubating3.jpg, Incubating in Copiapó, Chile Hummingbird-nest-2021.png, Nest with two eggs in
San Jose, California San Jose, officially San José (; ; ), is the largest city in Northern California Northern California (colloquially known as NorCal) is a geographic and cultural region that generally comprises the northern portion of the U.S. state of Califor ...
Hummingbird nest with two chicks in Santa Monica, CA. Photo taken June 26, 2006.jpg, Nest with two nestlings in
Santa Monica, California Santa Monica () is a beachfront city in western Los Angeles County, California, United States. Situated on Santa Monica Bay, it is bordered on five sides by different neighborhoods of the city of Los Angeles: Pacific Palisades, Los Angeles, Pac ...

Santa Monica, California
Calliope-nest edit.jpg, Feeding two nestlings in
Grand Teton National Park Grand Teton National Park is an American national park in northwestern Wyoming. At approximately , the park includes the major peaks of the Teton Range as well as most of the northern sections of the valley known as Jackson Hole. Grand Teton Na ...

Grand Teton National Park
AnnasHummingbird-NestCollage.png, Fallen
Anna's hummingbird Anna's hummingbird (''Calypte anna'') is a medium-sized bird species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecu ...

Anna's hummingbird
nest in
Ventura County, California Ventura County is a County (United States), county in Southern California, the southern part of the U.S. state of California. As of the 2020 United States Census, 2020 census, the population was 843,843. The largest city is Oxnard, California, O ...
, shown next to a
toothpick A toothpick is a small thin stick of wood Wood is a porous and fibrous structural tissue found in the stems and roots of tree In botany, a tree is a perennial plant with an elongated Plant stem, stem, or trunk (botany), trunk, ...

toothpick
for scale


Feather colors

To serve
courtship Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...

courtship
and territorial
competition Competition is a rivalry A rivalry is the state of two people or groups engaging in a lasting competitive relationship. Rivalry is the "against each other" spirit between two competing sides. The relationship itself may also be called "a ri ...
, many male hummingbirds have
plumage Plumage ( "feather") is a layer of feather Feathers are epidermal growths that form a distinctive outer covering, or plumage Plumage ( "feather") is a layer of feather Feathers are epidermal growths that form a distinctive outer ...
with bright, varied coloration resulting both from
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compound , CH4; is among the simplest organic compou ...
ation in the feathers and from
prism A prism An optical prism is a transparent optics, optical element with flat, polished surfaces that refraction, refract light. At least one surface must be angled—elements with two parallel surfaces are not prisms. The traditional geometrical ...

prism
al cells within the top layers of feathers of the head,
gorget A gorget , from the French ' meaning throat In vertebrate anatomy, the throat is the front part of the neck, positioned in front of the vertebra. It contains the Human pharynx, pharynx and larynx. An important section of it is the epiglotti ...
, breast, back and wings. When
sunlight Sunlight is a portion of the given off by the , in particular , , and light. On , sunlight is and through , and is obvious as when the Sun is above the . When direct is not blocked by s, it is experienced as sunshine, a combination of b ...

sunlight
hits these cells, it is split into
wavelength In physics Physics is the that studies , its , its and behavior through , and the related entities of and . "Physical science is that department of knowledge which relates to the order of nature, or, in other words, to the regular su ...

wavelength
s that reflect to the observer in varying degrees of intensity, with the feather structure acting as a
diffraction grating In optics, a diffraction grating is an optical component with a periodic structure that diffraction, diffracts light into several beams travelling in different directions (i.e., different diffraction angles). The emerging coloration is a form ...

diffraction grating
.
Iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to Gradient, gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, ...
hummingbird colors result from a combination of refraction and pigmentation, since the diffraction structures themselves are made of
melanin Melanin (; from el, μέλας ''melas'', "black, dark") is a broad term for a group of natural pigments found in most organisms. Melanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the ami ...

melanin
, a pigment, and may also be colored by
carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic Organic may refer to: * Organic, of or relating to an organism, a living entity * Organic, of or relating to an anatomical organ (anatomy), organ Chemistry * Organ ...
pigmentation and more subdued black, brown or gray colors dependent on melanin. By merely shifting position, feather regions of a muted-looking bird can instantly become fiery red or vivid green. In courtship displays for one example, males of the colorful
Anna's hummingbird Anna's hummingbird (''Calypte anna'') is a medium-sized bird species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecu ...

Anna's hummingbird
orient their bodies and feathers toward the sun to enhance the display value of iridescent plumage toward a female of interest. One study of Anna's hummingbirds found that dietary
protein Proteins are large biomolecule , showing alpha helices, represented by ribbons. This poten was the first to have its suckture solved by X-ray crystallography by Max Perutz and Sir John Cowdery Kendrew in 1958, for which they received a No ...

protein
was an influential factor in feather color, as birds receiving more protein grew significantly more colorful
crown '' File:서봉총 금관 금제드리개.jpg, The Seobongchong Golden Crown of Ancient Silla, which is 339th National Treasure of South Korea. It is basically following the standard type of Silla's Crown. It was excavated by Swedish Crown Pri ...
feathers than those fed a low-protein diet. Additionally, birds on a high-protein diet grew yellower (higher
hue In color theory In the visual arts The visual arts are art forms such as painting Painting is the practice of applying paint Paint is any pigmented liquid, liquefiable, or solid mastic composition that, after applica ...

hue
) green tail feathers than birds on a low-protein diet.


Aerodynamics of flight

Hummingbird flight has been studied intensively from an
aerodynamic study at Wallops Flight Facility, Wallops Island in 1990. A vortex is created by passage of an aircraft wing, revealed by smoke. Vortices are one of the many phenomena associated with the study of aerodynamics. Aerodynamics, from Greek language, ...
perspective using wind tunnels and high-speed
video camera A video camera is a camera A camera is an optical Optics is the branch of physics Physics (from grc, φυσική (ἐπιστήμη), physikḗ (epistḗmē), knowledge of nature, from ''phýsis'' 'nature'), , is the natural ...

video camera
s. Two studies of
rufous Rufous () is a color Color (American English American English (AmE, AE, AmEng, USEng, en-US), sometimes called United States English or U.S. English, is the set of varieties of the English language native to the United States. C ...
or
Anna's hummingbird Anna's hummingbird (''Calypte anna'') is a medium-sized bird species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecu ...

Anna's hummingbird
s in a wind tunnel used
particle image velocimetryParticle image velocimetry (PIV) is an optical method of flow visualization used in education and research. It is used to obtain instantaneous velocity The velocity of an object is the rate of change of its position with respect to a fram ...
techniques to investigate the lift generated on the bird's upstroke and downstroke. The birds produced 75% of their weight support during the downstroke and 25% during the upstroke, with the wings making a "figure 8" motion. Many earlier studies had assumed that
lift Lift or LIFT may refer to: Physical devices * Elevator, or lift, a device used for raising and lowering people or goods ** Rack lift, a type of elevator ** Ski lift, an aerial or surface lift for uphill transport ** Space elevator, a hypothetical ...
was generated equally during the two phases of the wingbeat cycle, as is the case of insects of a similar size. This finding shows that hummingbird is similar to, but distinct from, that of hovering insects such as the
hawk moth The Sphingidae are a family of Moth, moths (Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics ...

hawk moth
. Further studies using
electromyography Electromyography (EMG) is a technique for evaluating and recording the electrical activity produced by skeletal muscles.Kamen, Gary. Electromyographic Kinesiology. In Robertson, DGE et al. Research Methods in Biomechanics. Champaign, IL: Human K ...
in hovering rufous hummingbirds showed that
muscle strain A strain is an acute or Chronic condition, chronic soft tissue injury that occurs to a muscle, tendon, or both. The equivalent injury to a ligament is a sprain. Generally, the muscle or tendon overstretches and partially tears, under more physica ...
in the
pectoralis major The pectoralis major () is a thick, fan-shaped or triangular convergent muscle, situated at the chest of the human body. It makes up the bulk of the chest muscles Skeletal muscles (commonly referred to as muscles) are Organ (biology), organs ...

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. Because of their flying technique, these birds no longer have an
alula The alula , or bastard wing, (plural ''alulae'') is a small projection on the anterior edge of the wing of modern birds and a few non-avian dinosaurs. The word is Latin and means "winglet"; it is the diminutive of ''ala'', meaning "wing". The alu ...

alula
. The giant hummingbird's wings beat as few as 12/sec and the wings of typical hummingbirds beat up to 80/sec. 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. 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.


Feather sonation


Courtship dives

When courting, the male Anna's hummingbird ascends some above a female, before diving at a speed of , equal to 385 body lengths/sec – producing a high-pitched sound near the female at the
nadir The nadir (, ; ar, نظير, naẓīr, counterpart) is the direction pointing directly ''below'' a particular location; that is, it is one of two vertical direction In astronomy Astronomy (from el, ἀστρονομία, literally ...
of the dive. 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 falcon The peregrine falcon (''Falco peregrinus''), also known as the peregrine, and historically as the duck hawk in North America, is a Cosmopolitan distribution, cosmopolitan bird of prey (Bird of prey, raptor) in the family (biology), family Falco ...

peregrine falcon
s in pursuit of prey. At maximum descent speed, about 10 g of
gravitational force Gravity (), or gravitation, is a natural phenomenon Types of natural phenomena include: Weather, fog, thunder, tornadoes; biological processes, decomposition, germination seedlings, three days after germination. Germination is t ...

gravitational force
occur in the courting hummingbird during a dive (Note:
G-force The gravitational force equivalent, or, more commonly, g-force, is a measurement of the type of force per unit mass – typically acceleration – that causes a perception of weight In science Science () is a systematic ente ...

G-force
is generated as the bird pulls out of the dive). By comparison to humans, this is a G-force acceleration well beyond the threshold of causing near loss of
consciousness Consciousness, at its simplest, is sentience or awareness of internal and external existence. Despite millennia of analyses, definitions, explanations and debates by philosophers and scientists, consciousness remains puzzling and controversial ...

consciousness
in
fighter pilot A fighter pilot is a Military aviation, military aviator trained to engage in air-to-air combat, Air-to-ground weaponry, air-to-ground combat and sometimes Electronic-warfare aircraft, electronic warfare while in the cockpit of a fighter aircraf ...

fighter pilot
s (occurring at about +5 Gz) during flight of fixed-wing aircraft in a high-speed
banked turn A banked turn (or banking turn) is a turn or change of direction in which the vehicle banks or inclines, usually towards the inside of the turn. For a road or railroad this is usually due to the roadbed having a transverse down-slope towards the ...

banked turn
. The outer tail feathers of male Anna's (''Calypte anna'') and ''Selasphorus'' hummingbirds (e.g., Allen's, calliope) vibrate during
courtship Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...

courtship
display dives and produce an audible chirp caused by aeroelastic flutter. 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. The bird can sing at the same frequency as the tail-feather chirp, but its small syrinx is not capable of the same volume. The sound is caused by the
aerodynamics Aerodynamics, from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is appr ...
of rapid air flow past tail feathers, causing them to flutter in a
vibration Vibration is a mechanical phenomenon whereby oscillation Oscillation is the repetitive variation, typically in time Time is the indefinite continued sequence, progress of existence and event (philosophy), events that occur in an apparentl ...

vibration
, which produces the high-pitched sound of a courtship dive. 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 The calliope hummingbird (''Selasphorus calliope'') (pronunciation kə-ˈlī-ə-pē) is the smallest bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless bea ...
,
broad-tailed hummingbird The broad-tailed hummingbird (''Selasphorus platycercus'') is a medium-sized hummingbird Hummingbirds are birds native to the Americas and constitute the biological family Trochilidae. There are about 360 species. They occur from Alaska to ...
,
rufous hummingbird The rufous hummingbird (''Selasphorus rufus'') is a small hummingbird, about long with a long, straight and slender bill. These birds are known for their extraordinary flight skills, flying during their migratory transits. It is one of seven s ...
,
Allen's hummingbird Allen's hummingbird (''Selasphorus sasin'') is a species of hummingbird that breeds in the western United States. It is one of seven species in the genus ''Selasphorus''. Description Allen's hummingbird is a small bird, with mature adults reach ...

Allen's hummingbird
, and streamertail, as well as the tail of the
Costa's hummingbird Costa's hummingbird (''Calypte costae'') is a bird species in the hummingbird family Trochilidae. It breeds in the arid region of the southwest United States and northwest Mexico; it winters in western Mexico. Taxonomy Costa's hummingbird was spe ...
and the
black-chinned hummingbird The black-chinned hummingbird (''Archilochus alexandri'') is a small hummingbird occupying a broad range of habitats. It is migratory, spending winter as far south as Mexico Mexico ( es, México ; Nahuan languages: ), officially the United ...
, and a number of related species. The
harmonic A harmonic is any member of the harmonic series Harmonic series may refer to either of two related concepts: *Harmonic series (mathematics) *Harmonic series (music) {{Disambig .... The term is employed in various disciplines, including music ...
s of sounds during courtship dives vary across species of hummingbirds.


Wing feather trill

Male rufous and broad-tailed hummingbirds (genus ''
Selasphorus ''Selasphorus'' is a genus of hummingbirds from Middle America (Americas), Middle and North America. Taxonomy The genus ''Selasphorus'' was introduced in 1832 by the English naturalist William John Swainson to accommodate the rufous hummingbird w ...
'') have a distinctive wing feature during normal flight that sounds like jingling or a buzzing shrill whistle. 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. Behaviorally, the trill serves several purposes: * 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 Alaska (; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Central Alaskan Yup'ik language, Yup'ik: ''Alaskaq''; tli, Anáaski) is a U.S. state in the Western United States, on the northwest extremity of the country's West Coast of the United State ...

Alaska
to
Tierra del Fuego #REDIRECT Tierra del Fuego #REDIRECT Tierra del Fuego#REDIRECT Tierra del Fuego Tierra del Fuego (, ; Spanish for "Land of Fire", formerly also Fireland in English) is an archipelago An archipelago ( ), sometimes called an island group or i ...

Tierra del Fuego
, including the
Caribbean The Caribbean (, ; es, Caribe; french: Caraïbes; ht, Karayib; also gcf, label=Antillean Creole Antillean Creole (Antillean French Creole, Kreyol, Kwéyòl, Patois) is a French-based creole, which is primarily spoken in the Lesser Antilles ...
. 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 . The greatest
species richness File:Global Amphibian Richness Grids, 2015 Release, All Amphibians (28794889801).jpg, 300px, Global amphibian richness (2015) Species richness is the number of different species represented in an community (ecology), ecological community, landscape ...
is in humid tropical and subtropical forests of the northern Andes and adjacent foothills, but the number of species found in the
Atlantic Forest The Atlantic Forest ( pt, Mata Atlântica) is a South America South America is a continent A continent is any of several large landmasses. Generally identified by convention (norm), convention rather than any strict criteri ...

Atlantic Forest
, Central America or southern
Mexico Mexico, officially the United Mexican States, is a country A country is a distinct territorial body or political entity A polity is an identifiable political entity—any group of people who have a collective identity, who are organi ...

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 Chile, officially the Republic of Chile, is a country in the western part of South America South America is a entirely in the and mostly in the , with a relatively small portion in the . It can also be described as the southern ...

Chile
each,
Colombia Colombia ( , ; ), officially the Republic of Colombia, is a transcontinental country This is a list of countries located on more than one continent A continent is one of several large landmasses. Generally identified by conv ...

Colombia
alone has more than 160 and the comparably small
Ecuador Ecuador ( ; ; Quechua Quechua may refer to: *Quechua people, several indigenous ethnic groups in South America, especially in Peru *Quechuan languages, a Native South American language family spoken primarily in the Andes, derived from a ...

Ecuador
has about 130 species. The migratory
ruby-throated hummingbird The ruby-throated hummingbird (''Archilochus colubris'') is a species of hummingbird Hummingbirds are birds native to the Americas and constitute the biological family Trochilidae. There are about 360 species. They occur from Alaska to Tier ...

ruby-throated hummingbird
breeds in a range from the
Southeastern United States The southeastern United States, also referred to as the American Southeast or simply the Southeast, is broadly the eastern portion of the southern United States The Southern United States, also referred to as the Southern States, the Ame ...
to
Ontario ("Loyal she began, loyal she remains") , Label_map = yes , image_map = Ontario in Canada 2.svg , map_alt = Map showing Ontario's location east/central of Canada. , coordinates = , cap ...

Ontario
, while the
black-chinned hummingbird The black-chinned hummingbird (''Archilochus alexandri'') is a small hummingbird occupying a broad range of habitats. It is migratory, spending winter as far south as Mexico Mexico ( es, México ; Nahuan languages: ), officially the United ...
, its close relative and another migrant, is the most widespread and common species in the southwestern United States. The
rufous hummingbird The rufous hummingbird (''Selasphorus rufus'') is a small hummingbird, about long with a long, straight and slender bill. These birds are known for their extraordinary flight skills, flying during their migratory transits. It is one of seven s ...
is the most widespread species in western North America,Williamson, S. L. (2002). ''A Field Guide to Hummingbirds of North America'' (Peterson Field Guide Series). Houghton Mifflin Co., Boston. and the only hummingbird to be recorded outside of the Americas, having occurred in the
Chukchi Peninsula The Chukchi Peninsula (also Chukotka Peninsula or Chukotski Peninsula; russian: Чуко́тский полуо́стров, ''Chukotskiy poluostrov'', short form russian: Чуко́тка, ''Chukotka''), at about 66° N 172° W, is the eastern ...
of Russia.


Migration

Most North American hummingbirds migrate southward in fall to spend winter in Mexico, the Caribbean Islands, or Central America. 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. Among these are Anna's hummingbird, a common resident from southern Arizona and inland California, and the , a winter resident from Florida across the Gulf Coast to
South Texas South Texas is a region of the U.S. state of Texas Texas (, ; : ''Texas'', ''Tejas'') is a state in the region of the . At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest ...
. Ruby-throated hummingbirds are common along the
Atlantic flyway The Atlantic Flyway is a major north-south flyway for migratory birds in North America. The route generally starts in Greenland, then follows the Atlantic coast of Canada, then south down the Atlantic Coast south to the tropical areas of South A ...
, and migrate in summer from as far north as
Atlantic Canada Atlantic Canada, also called the Atlantic provinces, a term developed for the convenience of the federal government after Newfoundland Newfoundland and Labrador (, ) is the easternmost provinces and territories of Canada, province of Canada, ...

Atlantic Canada
, returning to
Mexico Mexico, officially the United Mexican States, is a country A country is a distinct territorial body or political entity A polity is an identifiable political entity—any group of people who have a collective identity, who are organi ...

Mexico
, South America, southern Texas, and
Florida Florida is a U.S. state, state located in the Southeastern United States, Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia (U.S. state), Geor ...

Florida
to winter. During winter in southern
Louisiana Louisiana (Standard French Standard French (in French: ''le français standard'', ''le français normé'', ''le français neutre'' eutral Frenchor ''le français international'' nternational French is an unofficial term for a standard ...

Louisiana
, black-chinned, buff-bellied, calliope, Allen's, Anna's, ruby-throated, rufous, broad-tailed, and broad-billed hummingbirds are present. The
rufous hummingbird The rufous hummingbird (''Selasphorus rufus'') is a small hummingbird, about long with a long, straight and slender bill. These birds are known for their extraordinary flight skills, flying during their migratory transits. It is one of seven s ...
breeds farther north than any other species of hummingbird, often breeding in large numbers in temperate North America and wintering in increasing numbers along the coasts of the subtropical
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin 400px, Diagrammatic cross-section of an ocean basin, showing the various geographic features In hydrology Hydrology (from Greek: wikt:ὕδωρ, ὕδωρ, "hýdōr" meaning ...

Gulf of Mexico
and Florida, rather than in western or central Mexico. By migrating in spring as far north as the
Yukon Yukon ( ; ; formerly called Yukon Territory and sometimes referred to as The Yukon) is the smallest and westernmost of Canada's three territories. It also is the least populated province or territory in Canada, with a population of 35,874 peo ...

Yukon
or southern
Alaska Alaska (; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Central Alaskan Yup'ik language, Yup'ik: ''Alaskaq''; tli, Anáaski) is a U.S. state in the Western United States, on the northwest extremity of the country's West Coast of the United State ...

Alaska
, 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. As calculated by
displacement Displacement may refer to: Physical sciences Mathematics and Physics *Displacement (geometry), is the difference between the final and initial position of a point trajectory (for instance, the center of mass of a moving object). The actual path c ...
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. By comparison, the 13-inch-long
Arctic tern#REDIRECT Arctic tern {{Redirect category shell, 1= {{R from move {{R from other capitalisation ...

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. The northward migration of rufous hummingbirds occurs along the
Pacific flyway 300px, Waterfowl flyways in the United States. The Pacific Flyway is in green. The Pacific Flyway is a major north-south flyway A flyway is a flight path used by large numbers of birds while bird migration, migrating between their breeding gr ...
and may be time-coordinated with flower and tree-leaf emergence in spring in early March, and also with availability of insects as food. Arrival at breeding grounds before nectar availability from mature flowers may jeopardize breeding opportunities.


Diet and specializations for food gathering

For
nutrition Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It includes ingestion, Absorption (biology), absorption, Assimilation (biology), assimilation, biosynthesis, catabolism and excretion. ...
, hummingbirds eat a variety of insects, including
mosquito Mosquitoes are members of a group of almost 3,600 species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defin ...

mosquito
es,
fruit flies Fruit fly may refer to: Organisms * Drosophilidae, a family of smaller flies, including: ** ''Drosophila'', the genus of small fruit flies and vinegar flies ** ''Drosophila melanogaster'' or common fruit fly, an important model organism in modern b ...

fruit flies
,
gnat GNAT is a free-software Free software (or libre software) is computer software Software is a collection of Instruction (computer science), instructions and data (computing), data that tell a computer how to work. This is in contrast t ...

gnat
s in flight, or
aphid Aphids are small sap-sucking insects and members of the Taxonomic rank, superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Erio ...

aphid
s on leaves and
spider Spiders (order Order or ORDER or Orders may refer to: * Orderliness Orderliness is associated with other qualities such as cleanliness Cleanliness is both the abstract state of being clean and free from germs, dirt, trash, or waste, and the ...

spider
s in their webs. The lower
beak The beak, bill, and/or rostrum is an external anatomical structure found mostly in birds Birds are a group of warm-blooded vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellu ...

beak
of hummingbirds is flexible and can bend as much as 25 degrees when it widens at the base, making a larger surface for catching insects. Hummingbirds hover within insect
swarm Swarm behaviour, or swarming, is a collective animal behaviour, collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or an ...
s in a method called "hover-
hawking Hawking may refer to: * Stephen Hawking (1942–2018), English theoretical physicist and cosmologist **Hawking (2004 film), ''Hawking'' (2004 film), about Stephen Hawking **Hawking (2013 film), ''Hawking'' (2013 film), about Stephen Hawking * Hawk ...
" to facilitate feeding. 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
nutrient A nutrient is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance, a material with a definite chemical composition * Matter, anything that has mass and t ...
s, requiring hummingbirds to meet their nutritional needs by consuming insects. 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. Hummingbirds digest their food rapidly due to their small size and high metabolism; a mean retention time less than an hour has been reported. 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 Starvation is a severe deficiency in caloric energy In physics Physics is the that studies , its , its and behavior through , and the related entities of and . "Physical science is that department of knowledge which relates t ...

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 The hippocampus (via Latin from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its popul ...

hippocampus
, a brain region facilitating spatial memory used to map flowers previously visited during nectar foraging. Hummingbird beaks are flexible and their shapes vary dramatically as an adaptation for specialized feeding. 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 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 Gesneriaceae, the gesneriad family, is a family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or soci ...
. 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). When catching insects in flight, a hummingbird's jaw bending, flexes downward to widen the beak, gape for successful capture.


Perception of sweet nectar

Perception of sweetness in nectar evolved in hummingbirds during their genetic divergence from insectivorous swifts, their closest bird relatives. Although the only known sweet sensory receptor, called TAS1R2, T1R2, is absent in birds, receptor expression studies showed that hummingbirds adapted a carbohydrate receptor from the TAS1R1, T1R1-TAS1R3, T1R3 receptor, identical to the one perceived as umami in humans, essentially repurposing it to function as a nectar sweetness receptor. 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.


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. 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.Mosher
"High-Speed Video Shows How Hummingbirds Really Drink"
Wired.com. May 2, 2011.
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. Consequently, tongue flexibility enables accessing, transporting and unloading nectar.


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. 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 Pollination is the transfer of pollen Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes (sperm cells). Pollen grains have a hard coat ...

pollination
their feeding naturally provides. sucrose, White granulated sugar is used in hummingbird feeders in a 25% concentration as a common recipe, although hummingbirds will defend feeders more aggressively when sugar content is at 35%, indicating preference for nectar with higher sugar content. Organic and "raw" sugars contain iron, which can be harmful, and brown sugar, agave syrup, molasses, and Sugar substitute, artificial sweeteners also should not be used. Honey is made by
bee Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...

bee
s 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. Allura Red AC, Red food dye was once thought to be a favorable ingredient for the nectar in home feeders, but it is unnecessary. 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. Although some commercial products contain small amounts of nutritional additives, hummingbirds obtain all necessary nutrients from the insects they eat, rendering added nutrients unnecessary.


Visual cues of foraging

Hummingbirds have exceptional visual acuity providing them with discrimination of food sources while foraging. 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. 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. In at least one hummingbird species – the green-backed firecrown (''Sephanoides sephaniodes'') – flower colors preferred are in the red-green
wavelength In physics Physics is the that studies , its , its and behavior through , and the related entities of and . "Physical science is that department of knowledge which relates to the order of nature, or, in other words, to the regular su ...

wavelength
for the bird's bird vision, visual system, providing a higher contrast (vision), contrast than for other flower colors. 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 (ethology), dominance when foraging among other hummingbird species with less colorful plumage. The ability to discriminate colors of flowers and plumage is enabled by a
visual system The visual system comprises the sensory organ A sense is a biological system A biological system is a complex network which connects several biologically relevant entities. Biological organization spans several scales and are determined b ...
having four single cone cells and a double cone screened by photoreceptor cell, photoreceptor oil droplets which enhance color discrimination.


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 olfaction, smell the presence in nectar of insect defensive chemicals (such as formic acid) and aggregation pheromones of foraging ants, which discourage feeding. Another study showed that hummingbirds are attracted to the nectar aroma of certain flowers by the combination of Volatile organic compound, volatile monoterpenes, including d-limonene, β-myrcene, and β-ocimene.


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. The Hemaris, hummingbird moth is often mistaken for a hummingbird.


In myth and culture

*Aztecs wore hummingbird amulet, talismans, both artistic representations of hummingbirds and fetishism, 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. * 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. The Nahuatl word ''huitzil'' (hummingbird) is an onomatopoeia, 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 people, Zuni cultures have a 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. * Trinidad and Tobago, known as "The land of the hummingbird," displays a hummingbird on that nation's coat of arms of Trinidad and Tobago, coat of arms, 1-cent coin and emblem of its 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 that included golden wings supposed to follow the movements of her arms. However, it accidentally got damaged during the dress rehearsal, and she appeared onstage with a broken, drooping left wing.


Gallery

File:Female Ruby-Throated Hummingbird perched in Hudson, Ohio.jpg, Female ruby-throated hummingbird perched in Hudson, Ohio File:Humming birds feeding at 1500fps.webm, Hummingbirds feeding at 1500 fps File:Hummingbird feeding.jpg, Hummingbird feeding from a flower in the University of California Botanical Garden File:Hummingbird headshot.jpg, Hummingbird with yellow pollen on its beak in the University of California Botanical Garden File:Eugenes-fulgens-001.jpg, Talamanca hummingbird File:Hummingbird.jpg,
Costa's hummingbird Costa's hummingbird (''Calypte costae'') is a bird species in the hummingbird family Trochilidae. It breeds in the arid region of the southwest United States and northwest Mexico; it winters in western Mexico. Taxonomy Costa's hummingbird was spe ...
File:Juvenile Annas Hummingbird at San Francisco Botanical Garden - Sarah Stierch - E.jpg, Juvenile Anna's hummingbird with tongue sticking out File:Hummingbird Calypte anna in ggp 15n.jpg, ''Calypte anna'' perched File:Hummingbird Chicks in Nest in Cactus in Mesa, Arizona.jpg, Hummingbird chicks in a nest in a cactus in Mesa, Arizona File:Hummingbird Adult in Nest in Cactus in Mesa, Arizona.jpg, Hummingbird adult in its nest in a cactus in Mesa, Arizona File:Female Anna's Hummingbird.JPG, A female Anna's hummingbird 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


References


External links


The Hummingbird Website
Hummingbird photos, videos, articles, links, frequently asked questions
High-resolution photo gallery of almost 100 species


* [https://web.archive.org/web/20170301202153/http://www.learner.org/jnorth/tm/humm/tongue_fluid_trap.html Video of hummingbird tongue acting as a micropump during nectar feeding] {{Authority control Hummingbirds, Birds of the Americas, * Trochiliformes, * Articles containing video clips Extant Rupelian first appearances National symbols of Trinidad and Tobago Natural history of the Americas Taxa named by Nicholas Aylward Vigors