Editing Hummingbird (section)

== Evolution ==
Hummingbirds split from other members of Apodiformes, the insectivorous swifts (family Apodidae) and [[treeswift]]s (family Hemiprocnidae), about 42 million years ago, probably in [[Eurasia]].<ref name="mcguire2014"/> Despite their current New World distribution, the earliest species of hummingbird occurred in the early [[Oligocene]] ([[Rupelian]] about 34–28 million years ago) of Europe, belonging to the genus ''Eurotrochilus,'' having similar morphology to modern hummingbirds.<ref name="Mayr2004"/><ref>{{Cite journal |last=Mayr |first=Gerald |date=2007-01-01 |title=New specimens of the early Oligocene Old World hummingbird Eurotrochilus inexpectatus |url=https://doi.org/10.1007/s10336-006-0108-y |journal=Journal of Ornithology |language=en |volume=148 |issue=1 |pages=105–111 |doi=10.1007/s10336-006-0108-y |bibcode=2007JOrni.148..105M |issn=2193-7206 |s2cid=11821178|url-access=subscription }}</ref><ref>{{Cite journal |last1=Bochenski |first1=Zygmunt |last2=Bochenski |first2=Zbigniew M. |date=2008-04-01 |title=An Old World hummingbird from the Oligocene: a new fossil from Polish Carpathians |url=https://doi.org/10.1007/s10336-007-0261-y |journal=Journal of Ornithology |language=en |volume=149 |issue=2 |pages=211–216 |doi=10.1007/s10336-007-0261-y |bibcode=2008JOrni.149..211B |issn=2193-7206 |s2cid=22193761|url-access=subscription }}</ref>

===Phylogeny===
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.<ref name="mcguire2014"/>

A map of the hummingbird family tree – reconstructed from analysis of 284 [[species]] – shows rapid diversification from 22 million years ago.<ref name="sd">{{Cite web |date=3 April 2014 |title=Hummingbirds' 22-million-year-old history of remarkable change is far from complete |url=https://www.sciencedaily.com/releases/2014/04/140403132207.htm |access-date=30 September 2014 |website=ScienceDaily}}</ref> Hummingbirds fall into nine main clades – the [[Florisuginae|topazes]], [[Phaethornithinae|hermits]], [[Polytminae|mangoes]], [[Heliantheini|brilliants]], [[Lesbiini|coquettes]], the giant hummingbird, [[Lampornithini|mountaingems]], [[Mellisugini|bees]], and [[Trochilini|emeralds]] – defining their relationship to [[nectar]]-bearing [[flowering plant]]s which attract hummingbirds into new geographic areas.<ref name="mcguire2014"/><ref name="mcguire2007"/><ref name="mcg08">{{Cite journal |last1=McGuire |first1=Jimmy A. |last2=Witt |first2=Christopher C. |last3=Remsen |first3=J.V. Jr. |last4=Dudley |first4=R. |last5=Altshuler |first5=Douglas L. |date=2008 |title=A higher-level taxonomy for hummingbirds |journal=Journal of Ornithology |volume=150 |issue=1 |pages=155–165 |doi=10.1007/s10336-008-0330-x |issn=0021-8375 |s2cid=1918245}}</ref>

[[Molecular phylogenetics|Molecular phylogenetic]] studies of the hummingbirds have shown that the family is composed of nine major clades.<ref name="mcguire2007">{{Cite journal |last1=McGuire |first1=J.A. |last2=Witt |first2=C.C. |last3=Altshuler |first3=D.L. |last4=Remsen |first4=J.V. |date=2007 |title=Phylogenetic systematics and biogeography of hummingbirds: Bayesian and maximum likelihood analyses of partitioned data and selection of an appropriate partitioning strategy |journal=Systematic Biology |volume=56 |issue=5 |pages=837–856 |doi=10.1080/10635150701656360 |pmid=17934998 |doi-access=free}}</ref><ref name="mcguire2014">{{Cite journal |last1=McGuire |first1=J. |last2=Witt |first2=C. |last3=Remsen |first3=J.V. |last4=Corl |first4=A. |last5=Rabosky |first5=D. |last6=Altshuler |first6=D. |last7=Dudley |first7=R. |date=2014 |title=Molecular phylogenetics and the diversification of hummingbirds |journal=Current Biology |volume=24 |issue=8 |pages=910–916 |doi=10.1016/j.cub.2014.03.016 |pmid=24704078 |doi-access=free|bibcode=2014CBio...24..910M }}</ref> When [[Edward C. Dickinson|Edward Dickinson]] and [[James Van Remsen Jr.]] updated the ''[[Howard and Moore Complete Checklist of the Birds of the World]]'' for the 4th edition in 2013, they divided the hummingbirds into six subfamilies.<ref name="h&m4">{{Cite book |title=The Howard & Moore Complete Checklist of the Birds of the World |publisher=Aves Press |year=2013 |isbn=978-0-9568611-0-8 |editor-last=Dickinson |editor-first=E.C. |editor-link=Edward C. Dickinson |edition=4th |volume=1: Non-passerines |location=Eastbourne, UK |pages=105–136 |editor-last2=Remsen |editor-first2=J.V. Jr. |editor-link2=James Van Remsen Jr.}}</ref>

Molecular phylogenetic studies determined the relationships between the major groups of hummingbirds.<ref name=mcguire2014/><ref name=mcg08/> In the [[cladogram]] below, the English names are those introduced in 1997.<ref>{{Cite journal |last1=Bleiweiss |first1=R. |last2=Kirsch |first2=J.A. |last3=Matheus |first3=J.C. |date=1997 |title=DNA hybridization evidence for the principal lineages of hummingbirds (Aves:Trochilidae). |journal=Molecular Biology and Evolution |volume=14 |issue=3 |pages=325–343 |doi=10.1093/oxfordjournals.molbev.a025767 |pmid=9066799 |doi-access=free}}</ref> The [[scientific names]] are those introduced in 2013.{{Sfn|Dickinson|Remsen|2013|pp=105–136}}

{{Clade |style=font-size:100%;line-height:100%
|label1='''Trochilidae'''
|1={{Clade
   |1={{Clade
      |1=[[Florisuginae]] – topazes
      |2=[[Phaethornithinae]] – hermits
      }}
   |2={{Clade
      |1=[[Polytminae]] – mangoes
      |2={{Clade
         |label1=[[Lesbiinae]]
         |1={{Clade
            |1=[[Heliantheini]] – brilliants
            |2=[[Lesbiini]] – coquettes
            }}
         |2={{Clade
            |1=[[Patagoninae]] – giants
            |label2=[[Trochilinae]]
            |2={{Clade
               |1={{Clade
                  |1=[[Lampornithini]] – mountain gems
                  |2=[[Mellisugini]] – bees
                  }}
               |2=[[Trochilini]] – emeralds
               }}
            }}
         }}
      }}
   }}
}}

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]]ary pattern, as many as 140 hummingbird species can coexist in a specific region, such as the Andes [[Mountain range|range]].<ref name="sd"/>

The hummingbird [[Phylogenetic tree|evolutionary tree]] shows that one key evolutionary factor appears to have been an altered [[taste receptor]] that enabled hummingbirds to seek nectar.<ref>{{Cite journal |last1=Baldwin |first1=M.W. |last2=Toda |first2=Y. |last3=Nakagita |first3=T. |last4=O'Connell |first4=M.J. |last5=Klasing |first5=K.C. |last6=Misaka |first6=T. |last7=Edwards |first7=S.V. |last8=Liberles |first8=S. D. |year=2014 |title=Evolution of sweet taste perception in hummingbirds by transformation of the ancestral umami receptor |journal=Science |volume=345 |issue=6199 |pages=929–933 |bibcode=2014Sci...345..929B |doi=10.1126/science.1255097 |pmc=4302410 |pmid=25146290}}</ref>

Upon maturity, males of a particular species, ''Phaethornis longirostris,'' the [[long-billed hermit]], appear to be evolving a [[dagger]]-like weapon on the beak tip as a secondary [[sexual dimorphism|sexual trait]] to defend [[lek mating|mating areas]].<ref name="Rico-Guevara2015">{{Cite journal |author1=Rico-Guevara, A. |author2=Araya-Salas, M. |year=2015 |title=Bills as daggers? A test for sexually dimorphic weapons in a lekking hummingbird |journal=Behavioral Ecology |volume=26 |issue=1 |pages=21–29 |doi=10.1093/beheco/aru182|doi-access=free }}</ref>

===Geographic diversification===
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.<ref name="sd"/> Hummingbirds remain in dynamic diversification inhabiting ecological regions across South America, North America, and the Caribbean, indicating an enlarging [[evolutionary radiation]].<ref name="sd"/>

Within the same geographic region, hummingbird clades coevolved with nectar-bearing plant clades, affecting mechanisms of [[pollination]].<ref>{{Cite journal |author1=Abrahamczyk, S. |author2=Renner, S.S. |year=2015 |title=The temporal build-up of hummingbird/plant mutualisms in North America and temperate South America |journal=BMC Evolutionary Biology |volume=15 |issue=1 |page=104 |doi=10.1186/s12862-015-0388-z |pmc=4460853 |pmid=26058608 |doi-access=free |bibcode=2015BMCEE..15..104A }}</ref><ref>{{Cite journal |author1=Abrahamczyk, S. |author2=Souto-Vilarós, D. |author3=McGuire, J.A. |author4=Renner, S.S. |year=2015 |title=Diversity and clade ages of West Indian hummingbirds and the largest plant clades dependent on them: a 5–9 Myr young mutualistic system |url=https://zenodo.org/record/890511 |journal=Biological Journal of the Linnean Society |volume=114 |issue=4 |pages=848–859 |doi=10.1111/bij.12476}}</ref> 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'').<ref>{{Cite journal |last1=Abrahamczyk |first1=S. |last2=Souto-Vilaros |first2=D. |last3=Renner |first3=S. S. |year=2014 |title=Escape from extreme specialization: Passionflowers, bats and the sword-billed hummingbird |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1795 |pages=20140888 |doi=10.1098/rspb.2014.0888 |pmc=4213610 |pmid=25274372}}</ref>

=== Coevolution with ornithophilous flowers ===
[[File:Purple-throated carib hummingbird feeding.jpg|thumb|[[Purple-throated carib]] feeding at a flower]]

Hummingbirds are specialized [[nectarivore]]s tied to the [[ornithophily|ornithophilous]] flowers upon which they feed.<ref>{{Cite journal |last=Stiles |first=Gary |year=1981 |title=Geographical aspects of bird flower coevolution, with particular reference to Central America |url=https://www.biodiversitylibrary.org/pdf2/002816500087380.pdf |journal=Annals of the Missouri Botanical Garden |volume=68 |issue=2 |pages=323–351 |doi=10.2307/2398801 |jstor=2398801 |bibcode=1981AnMBG..68..323S |s2cid=87692272}}</ref> This [[coevolution]] implies that morphological traits of hummingbirds, such as bill length, bill curvature, and body mass, are correlated with morphological traits of plants, such as [[petal|corolla]] length, curvature, and volume.<ref>{{Cite journal |last1=Maglianesi |first1=M.A. |last2=Blüthgen |first2=N. |last3=Böhning-Gaese |first3=K. |last4=Schleuning |first4=M. |name-list-style=amp |date=2014 |title=Morphological traits determine specialization and resource use in plant–hummingbird networks in the Neotropics |url=https://www.researchgate.net/publication/268518487 |journal=Ecology |volume=95 |issue=12 |pages=3325–334 |doi=10.1890/13-2261.1|bibcode=2014Ecol...95.3325M }}</ref> Some species, especially those with unusual bill shapes, such as the sword-billed hummingbird and the [[eutoxeres|sicklebills]], are coevolved with a small number of flower species. Even in the most specialized hummingbird–plant mutualisms, the number of food plant lineages of the individual hummingbird species increases with time.<ref>{{Cite journal |last1=Abrahamczyk |first1=Stefan |last2=Poretschkin |first2=Constantin |last3=Renner |first3=Susanne S. |name-list-style=amp |date=2017 |title=Evolutionary flexibility in five hummingbird/plant mutualistic systems: testing temporal and geographic matching |journal=Journal of Biogeography |volume=44 |issue=8 |pages=1847–855 |doi=10.1111/jbi.12962 |bibcode=2017JBiog..44.1847A |s2cid=90399556}}</ref> The bee hummingbird (''Mellisuga helenae'') &ndash; the world's smallest bird &ndash; evolved to [[dwarfism]] likely because it had to compete with long-billed hummingbirds having an advantage for nectar foraging from specialized flowers, consequently leading the bee hummingbird to more successfully compete for flower foraging against insects.<ref>{{Cite magazine |last=Simon, Matt |date=10 July 2015 |title=Absurd Creature of the Week: The World's Tiniest Bird Weighs Less Than a Dime |url=https://www.wired.com/2015/07/absurd-creature-of-the-week-bee-hummingbird |magazine=Wired |access-date=8 March 2017}}</ref><ref>{{Cite journal|display-authors=3 |last1=Dalsgaard |first1=Bo |last2=Martín González |first2=Ana M. |last3=Olesen |first3=Jens M. |last4=Ollerton |first4=J |last5=Timmermann |first5=A |last6=Andersen |first6=L. H. |last7=Tossas |first7=A. G. |name-list-style=amp |year=2009 |title=Plant-hummingbird interactions in the West Indies: Floral specialisation gradients associated with environment and hummingbird size |journal=Oecologia |volume=159 |issue=4 |pages=757–766 |bibcode=2009Oecol.159..757D |doi=10.1007/s00442-008-1255-z |pmid=19132403 |s2cid=35922888}}</ref>

[[File:Colibri-thalassinus-001-edit.jpg|thumb|[[Lesser violetear]] at a flower]]

Many plants pollinated by hummingbirds produce flowers in shades of red, orange, and bright pink, although the birds take nectar from flowers of other colors. Hummingbirds can see [[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]] may render hummingbird-pollinated flowers relatively inconspicuous to most insects, thereby reducing [[nectar robbing]].<ref>{{Cite journal |last1=Rodríguez-Gironés |first1=M.A. |last2=Santamaría |first2=L. |name-list-style=amp |year=2004 |title=Why are so many bird flowers red? |journal=PLOS Biology |volume=2 |issue=10 |page=e350 |doi=10.1371/journal.pbio.0020350 |pmc=521733 |pmid=15486585 |doi-access=free }}</ref><ref>{{Cite journal |last=Altschuler |first=D. L. |year=2003 |title=Flower color, hummingbird pollination, and habitat irradiance in four Neotropical forests |journal=[[Biotropica]] |volume=35 |issue=3 |pages=344–355 |doi=10.1646/02113 |s2cid=55929111}}</ref> Hummingbird-pollinated flowers also produce relatively weak nectar (averaging 25% sugars) containing a high proportion of [[sucrose]], whereas insect-pollinated flowers typically produce more concentrated nectars dominated by [[fructose]] and [[glucose]].<ref>{{Cite journal |last1=Nicolson |first1=S.W. |last2=Fleming |first2=P.A. |name-list-style=amp |year=2003 |title=Nectar as food for birds: the physiological consequences of drinking dilute sugar solutions |url=http://researchrepository.murdoch.edu.au/id/eprint/4725 |journal=Plant Systematics and Evolution |volume=238 |issue=1–4 |pages=139–153 |doi=10.1007/s00606-003-0276-7 |bibcode=2003PSyEv.238..139N |s2cid=23401164|url-access=subscription }}</ref>

Hummingbirds and the plants they visit for nectar have a tight coevolutionary association, generally called a plant–bird [[mutualism (biology)|mutualistic network]].<ref name="Junker">{{Cite journal |last1=Junker |first1=Robert R. |last2=Blüthgen |first2=Nico |last3=Brehm |first3=Tanja |last4=Binkenstein |first4=Julia |last5=Paulus |first5=Justina |last6=Martin Schaefer |first6=H. |last7=Stang |first7=Martina |name-list-style=amp |date=2012-12-13 |title=Specialization on traits as basis for the niche-breadth of flower visitors and as structuring mechanism of ecological networks |journal=Functional Ecology |volume=27 |issue=2 |pages=329–341 |doi=10.1111/1365-2435.12005 |doi-access=free}}</ref> These birds show high specialization and modularity, especially in communities with high species richness. These associations are also observed when closely related hummingbirds, such as two species of the same genus, visit distinct sets of flowering species.<ref name=Junker/><ref name="Martín">{{Cite journal |last1=Martín González |first1=Ana M. |last2=Dalsgaard |first2=Bo |display-authors=etal |date=2015-07-30 |title=The macroecology of phylogenetically structured hummingbird-plant networks |journal=Global Ecology and Biogeography |volume=24 |issue=11 |pages=1212–224 |doi=10.1111/geb.12355 |bibcode=2015GloEB..24.1212M |hdl-access=free |hdl=10026.1/3407}}</ref>