Hover (behaviour)
{{Short description|Ability of some flying animals and insects}}
File:Eichlinia cucurbitae-hovering nectaring.jpg hovering]]
Hovering is the ability exhibited by some winged animals to remain relatively stationary in midair. Usually this involves rapid downward thrusts of the wings to generate upward lift. Sometimes hovering is maintained by flapping or soaring into a headwind; this form of hovering is called "wind hovering", "windhovering", or "kiting".
{{cite web
|url=https://web.stanford.edu/group/stanfordbirds/text/essays/Hovering_Flight.html
|title=Hovering Flight
|last1=Ehrlich
|first1=Paul R.
|last2=Dobkin
|first2=David S.
|last3=Wheye
|first3=Darryl
|publisher=stanfordbirds
}}
|last=Lekwa
|first=Steve
|date=Feb 22, 2021
|url=https://www.amestrib.com/story/opinion/2021/02/22/steve-lekwa-kiting-common-activity-iowas-raptors/4548379001/
|title=Kiting is a common activity for Iowa's raptors
|newspaper=Ames Tribune
}}
True hoverers
=Hummingbirds=
Hummingbirds hover over flowers to obtain nectar, flapping their wings at up to 70 beats per second.
{{cite web
|url=https://www.audubon.org/news/the-hummingbird-wing-beat-challenge
|title=The Hummingbird Wing Beat Challenge
|date=April 22, 2020
|publisher=National Audubon Society
}}
Archilochus colubris-male hovering.jpg|Ruby-throated hummingbird
Calypte costae-male hovering nectaring.jpg|Costa's hummingbird
Sword-billed hummingbird (male) at Guango Lodge, Ecuador (21310837273).jpg|Sword-billed hummingbird
=Bats=
Like hummingbirds, fruit bats and nectar bats hover over flowers while feeding on fruits or nectar. Comparison between bats and hummingbirds has revealed that these animals exert similar amounts of energy relative to body weight during hovering: hummingbirds can twist their wings more easily and are more aerodynamic, but bats have bigger wings and larger strokes.
{{cite web
|url=https://news.stanford.edu/2018/09/26/comparing-hovering-bats-hummingbirds/
|title=Stanford engineers study hovering bats and hummingbirds in Costa Rica
|date=September 26, 2018
|publisher=Stanford news
}}
{{cite news
|url=https://www.sciencenews.org/article/how-nectar-bats-fly-nowhere
|title=How nectar bats fly nowhere
|author=Susan Milius
|date=October 15, 2018
|publisher=Science News
}}
Choeronycteris mexicana, Mexican long-tongued bat (7371567444) 2.jpg|Mexican long-tongued bat
Lonchophylla robusta (on feeder).jpg|Orange nectar bat
=Kingfishers=
Small Kingfishers such as Belted kingfisher may hover over water before diving in to catch fish.
{{cite web
|url=https://www.audubon.org/field-guide/bird/belted-kingfisher
|title=Belted Kingfisher Megaceryle alcyon
|publisher=National Audubon Society
}}
Larger species such as Ringed kingfisher are too heavy to hover for more than a few seconds.
{{cite web
|url=https://www.allaboutbirds.org/guide/Ringed_Kingfisher/overview
|title=Ringed Kingfisher Megaceryle torquata
|publisher=Cornell University
}}
Megaceryle alcyon-male hovering.jpg|Belted kingfisher
Pied Kingfisher in hovering in Flight.jpg|Pied kingfisher
=Moths=
==Sphinx moths==
Some sphinx moths (family Sphingidae) are known as hummingbird moths for their ability to hover over flowers while nectaring. Moths are relatively heavy insects and sometimes hang on to the flower with their forelegs as they hover.
{{cite web
|url=https://www.massaudubon.org/nature-wildlife/insects-arachnids/hummingbird-moth
|title=Hummingbird Moth (Clearwing Moth)
|publisher=Mass Audubon
}}
Hemaris thysbe-hovering nectaring.jpg|Hummingbird clearwing
Macroglossum stellatarum-hovering nectaring.jpg|Hummingbird hawk-moth
Broad-bordered bee hawk-moth patuxent research refuge 7.9.21 DSC 1713.jpg|Broad-bordered bee hawk-moth
==Clearwing moths==
Some clearwing moths (family Sesiidae) also hover while nectaring
|last1=BROWN
|first1=LARRY N. and
|last2=MIZELL, III
|first2=RUSSELL F.
|date=1993
|journal=Tropical Lepidoptera
|title=THE CLEARWING BORERS OF FLORIDA (LEPIDOPTERA: SESIIDAE)
|url=https://journals.flvc.org/troplep/article/download/89327/85680/116170
|pages=1–21
|volume=4
|issue=3
|publisher=Florida Online Journals
}}
|last1=Potter
|first1=Daniel
|chapter=Clearwing Moths (Lepidoptera: Sesiidae)
|editor1-last=Capinera
|editor1-first=J.L.
|date=2 October 2014
|title=Encyclopedia of Entomology
|pages=928
|chapter-url=https://link.springer.com/referenceworkentry/10.1007/978-1-4020-6359-6_5123
|doi=10.1007/978-1-4020-6359-6_5123
|publisher=Springer, Dordrecht
|isbn=978-1-4020-6242-1
}}
or even puddling.
|url=https://www.inaturalist.org/observations/170969282
|title=Melittia celebica
|date=4 July 2023
|publisher=iNaturalist
}}
Females may also hover to inspect ovipositing sites.
|url=https://www.inaturalist.org/observations/118200645
|title=Pyropteron icteropus
|date=22 May 2022
|publisher=iNaturalist
}}
Chamaesphecia bibioniformis-male hovering.jpg|Chamaesphecia bibioniformis
Eichlinia calabaza-hovering.jpg|Southwestern squash vine borer
Eichlinia cucurbitae-hovering dorsal.jpg|Squash vine borer
File:Bumblebee mimic.webm|thumbtime=0|Hemaris diffinis is an excellent bumblebee mimic
=Hoverflies=
Hoverflies are flies that often hover over the plants they visit. This hovering behaviour is unlike that of hummingbirds since they do not feed in midair. Hovering in general may be a means of finding a food source; in addition, male hovering is often a territorial display seeking females,{{cite journal |last1=Collett |first1=T.S. |last2=Land |first2=M.F. |date=September 1978 |journal=Journal of Comparative Physiology |title=How hoverflies compute interception courses |url=https://www.scopus.com/record/display.uri?eid=2-s2.0-0001324077&origin=inward&txGid=ad71bb12cbc836c7415c848ba45e1ee5 |pages=191–204 |volume=125 |issue=3 |doi=10.1007/BF00656597 |publisher=Springer-Verlag |s2cid=26039329 |url-access=subscription }}
while female hovering serves to inspect ovipositing sites.
|last1=Almohamad
|first1=Raki
|last2=Verheggen
|first2=François J.
|last3=HaubrugeUniv
|first3=Éric
|date=2009
|journal=Biotechnologie, Agronomie, Société et Environnement
|title=Searching and oviposition behavior of aphidophagous hoverflies (Diptera: Syrphidae): a review
|url=https://www.researchgate.net/publication/26849775
|pages=467–481
|volume=13
|issue=3
}}
|url=https://ucanr.edu/sites/tuolumne_county_master_gardeners/files/147388.pdf
|title=Hover Flies, a Gardener's Friend
|author=Vera Strader
|publisher=University of California Agriculture and Natural Resources
}}
|url=https://bugguide.net/node/view/2317562
|title=Allograpta exotica ovipositing - Allograpta exotica - Female
|author=Peter Chen
|date=December 6, 2023
|publisher=Iowa State University
|website=Bugguide }}
Eupeodes_fumipennis-female_hovering.jpg|Western aphideater
Eupeodes_volucris-female_hovering.jpg|Large-tailed aphideater
Helophilus_fasciatus-female_hovering.jpg|Narrow-headed marsh fly
Syritta_pipiens-male_hovering.jpg|Thick-legged hoverfly
=Bee flies=
Bee flies are parasitoids that can dart about in the air with great agility. Males hover as a courtship display,
|url=https://www.nhm.ac.uk/resources-rx/files/bee-fly-identification-129900.pdf
|title=Bee-flies Bombylius sp.
|publisher=Natural History Museum
}}
|last1=Ferguson
|first1=David J
|last2=Yeates
|first2=David K
|url=https://www.researchgate.net/publication/288683339
|title=The courtship behavior of the bee fly Meomyia vetusta Walker (Diptera: Bombyliidae)
|date=May 2013
|journal=Australian Entomologist
|volume=40
|pages=89–92
|via=ResearchGate
}}
while females hover over ovipositing sites - usually the entrance of a host insect nest - and shoot eggs into the nest using an ejecting movement of their abdomen.
|last1=Boesi
|first1=Roberto
|last2=Polidori
|first2=Carlo
|last3=Andrietti
|first3=Francesco
|url=https://www.researchgate.net/publication/230785533
|title=Searching for the Right Target: Oviposition and Feeding Behavior in Bombylius Bee Flies (Diptera: Bombyliidae)
|date=March 2009
|journal=Zoological Studies
|volume=48
|issue=2
|pages=141–150
|via=ResearchGate
}}
Species that have a long proboscis can hover over flowers while feeding, much as hummingbirds do, though these flies may touch the flower with their legs for balance while hovering.
|url=https://www.buzzaboutbees.net/bee-fly.html
|title=The Large Bee Fly - Bombylius major
|date=April 20, 2021
|publisher=Buzzaboutbees
}}
Bombylius egg ejection 290310 1504.gif|Bombylius egg ejection
Bombylius major-hovering.jpg|Large bee-fly
Bombylius canescens-male hovering.jpg|Western bee-fly
Bombylella atra-hovering.jpg|Bombylella atra
Heterostylum robustum-hovering.jpg|Bomber fly
=Odonata=
Odonata is an insect order that includes dragonflies and damselflies. They are strong aviators renowned for their acrobatic flights, including the ability to hover, usually for a short pause during their ceaseless territorial patrols.
{{cite web
|url=https://australian.museum/learn/animals/insects/dragonflies-and-damselflies-order-odonata/
|title=Dragonflies and damselflies - Order Odonata
|author=David Britton
|date=Mar 7, 2023
|publisher=Australian Museum
}}
==Dragonflies==
In addition to short hovers while cruising, female dragonflies may hover over the water before or during oviposition, males may also hover-guard their mate at this time.
|last1=McMillan
|first1=Victoria E.
|date=June 1991
|journal=Animal Behaviour
|url=https://www.sciencedirect.com/science/article/abs/pii/S0003347205806360
|title=Variable mate-guarding behaviour in the dragonfly Plathemis lydia (Odonata: Libellulidae)
|pages=979–987
|volume=41
|issue=6
|doi=10.1016/S0003-3472(05)80636-0
|publisher=Elsevier B.V.
|s2cid=54360417
|url-access=subscription
}}
Aeshna juncea hovering.jpg|Sedge darner male
Anax junius-hovering.jpg|Green darner male
Plathemis lydia-female hovering.jpg|Common whitetail female
Tramea lacerata-flying tandem separated.jpg|Black saddlebags pair hovering over oviposition site
Tramea lacerata-guarded ovipositing.jpg|Black saddlebags male hover-guarding his mate
Pachydiplax longipennis-female ovipositing.webm|Blue dasher
Plathemis lydia-ovipositing.webm|Common whitetail
==Damselflies==
Some male damselflies hover in front of females or over the oviposition site during courtship; sometimes females also hover in response.
{{cite web
|url=https://uwm.edu/field-station/bug-of-the-week/river-jewelwing-damselfly/
|title=River Jewelwing Damselfly
|author=The BugLady
|date=December 14, 2022
|publisher=University of Wisconsin-Milwaukee
}}
|last1=Guillermo-Ferreiraa
|first1=Rhainer
|last2=Neissc
|first2=Ulisses Gaspar
|last3=Hamadad
|first3=Neusa
|last4=Bispo
|first4=Pitágoras C.
|date=2 October 2014
|journal=International Journal of Odonatology
|title=Behavior of the Amazonian damselfly Chalcopteryx scintillans McLachlan (Zygoptera: Polythoridae) and comments on its morphological distinction from C. rutilans (Rambur)
|url=https://worlddragonfly.org/article/13887890-2014-983189/
|pages=251–258
|volume=17
|issue=4
|doi=10.1080/13887890.2014.983189
|publisher=Worldwide Dragonfly Association
|doi-access=free
|bibcode=2014IJOdo..17..251G
}}
After mating, males may hover-guard their mate by either circling over her or by hovering while attached to her in tandem. Males hover-guarding in tandem do not need wings at all to remain suspended in the air; they are held aloft by clasping their mate with their abdomen, and can maintain their position even when the head and thorax are removed by predators.
{{cite web
|url=https://fieldguides.fieldmuseum.org/sites/default/files/rapid-color-guides-pdfs/388_0.pdf
|title=Damselflies of Chicagoland A Photo Field Guide, version 2
|author=Marla Garrison
|date=2011
|publisher=The Field Museum, Chicago
}}
{{cite web
|url=https://www.lakechamplaincommittee.org/learn/news/item/nature-note-dragonflies-and-damselflies
|title=Nature Note: Dragonflies and Damselflies
|date=August 31, 2009
|publisher=Lake Champlain Committee
}}
Calopteryx maculata-male hovering.jpg|Ebony jewelwing male hover-patrolling
Argia moesta-male hovering.jpg|Powdered dancer male hover-patrolling
Argia moesta-ovipositing.jpg|Powdered dancer males hover-guarding
Argia moesta-truncated tandem.jpg|Powdered dancer male posthumous hover-guarding
=Hymenoptera=
==Bees==
Many bee species, such as bumblebees, hover momentarily as they approach flowers to feed.
|url=https://www.buzzaboutbees.net/do-bumble-bees-hover.html
|title=Do Bumble Bees Hover?
|publisher=Buzzaboutbees
}}
Males of some species, including carder bees, hover while patrolling their territories.
|url=https://bestbeebrothers.com/blogs/blog/why-carpenter-bees-hover
|title=Why Do Carpenter Bees Hover?
|date=January 12, 2023
|publisher=Best Bee Brothers
}}
|url=https://entomology.ca.uky.edu/ef611
|title=Carpenter Bees
|author=MICHAEL F. POTTER
|publisher=University of Kentucky
}}
|url=https://entnemdept.ufl.edu/creatures/MISC/BEES/Anthidium_manicatum.html
|title=common name: European wool carder bee
|author1=Samantha Gallagher
|author2=Andrea Lucky
|date=December 2019
|publisher=University of Florida
}}
Bombus fervidus-hovering.jpg|Golden northern bumble bee
Xylocopa nasalis-hovering.jpg|Oriental carpenter bee
Anthidium manicatum-male hovering.jpg|European wool carder bee
Large eyed male carpenter bees primarily hover to protect their territory and attract female carpenter bees. Hovering allows them to spot intruders and other male competitors. When a male carpenter bee encounters an intruder, including a person and other mammals, it may dart towards it to chase it away.{{cite web|url=https://www.whatsthatbug.com/why-do-carpenter-bees-hover|title=Why Do Carpenter Bees Hover? Uncovering Their Mysterious Behavior
|work=whatsthatbug.com|date=13 October 2023 }}
File:CarpenderBeeHovering 1 prob4 63.webm hovering]]
==Wasps==
Among the social wasps, Stenogastrinae are known as hover wasps due to their distinctive hovering flight.
|last1=Carpenter
|first1=James Michael
|last2=Nguyen
|first2=Lien Thi Phuong
|url=https://www.researchgate.net/publication/227616275
|title=Keys to the genera of social wasps of South-East Asia (Hymenoptera: Vespidae)
|date=September 2003
|journal=Entomological Science
|volume=6
|issue=3
|pages=183–192
|publisher=American Museum of Natural History
|via=ResearchGate
|doi=10.1046/j.1343-8786.2003.00016.x
|s2cid=86823797
}}
Males often hover to display banding patterns on their abdomen as a territorial display.
|last1=BEANI
|first1=L.
|last2=TURILLAZZI
|first2=S.
|url=https://www.sciencedirect.com/science/article/abs/pii/S0003347299911016
|title=Stripes display in hover-wasps (Vespidae: Stenogastrinae): a socially costly status badge
|date=June 1999
|journal=Animal Behaviour
|volume=57
|issue=6
|pages=1233–1239
|publisher=The Association for the Study of Animal Behaviour
|via=Elsevier
|doi=10.1006/anbe.1999.1101
|pmid=10373256
|s2cid=45496129
|url-access=subscription
}}
|last1=Turillazzi
|first1=S.
|last2=Francescato
|first2=E.
|url=https://www.sciencedirect.com/science/article/abs/pii/S0003347299911016
|title=Patrolling behaviour and related secretory structures in the males of some Stenogastrine wasps
|date=1990
|journal=Insectes Sociaux
|volume=37
|pages=146–157
|via=Springer Nature
|doi=10.1007/BF02224027
|s2cid=25746201
|url-access=subscription
}}
Among the solitary wasps, parasitoid species such as scoliid wasps exhibit hovering behaviour while hunting for prey to feed their larvae.
|url=https://www.dailypress.com/1990/08/26/flying-pests-are-more-than-just-annoying/
|title=FLYING PESTS ARE MORE THAN JUST ANNOYING
|date=July 30, 2019
|publisher=Daily Press
}}
|url=https://extension.umd.edu/resource/scoliid-wasps-lawns/
|title=Scoliid Wasps - Lawns
|date=March 1, 2023
|publisher=University of Maryland Extension
}}
Males of some parasitoids may hover briefly while they patrol their territories, seeking females and chasing away rivals.
|url=https://mdc.mo.gov/discover-nature/field-guide/eastern-cicada-killer-wasp
|title=Eastern Cicada-Killer Wasp
|publisher=Missouri Department of Conservation
}}
|url=https://content.ces.ncsu.edu/cicada-killer-wasp
|title=Cicada Killer Wasp
|publisher=N.C. Cooperative Extension
}}
|url=https://www.ydr.com/story/life/2015/07/13/masterful-gardening-sand-wasps-our-allies-pest-control/72142742/
|title=Masterful Gardening: Sand wasps - our allies in pest control
|author=Connie Schmotzer
|date=July 13, 2015
|publisher=YorkDailyRecord
}}
Sphecius speciosus-male hovering.jpg|Eastern cicada killer hover patrol
Wind hoverers
=Raptors=
Many birds of prey such as kestrels, harriers, and members of the Buteo genus can "windhover" by facing the wind.
{{cite web
|url=https://birdfact.com/articles/kestrel-hovering
|title=Kestrel Hovering: A Complete Guide
|date=March 15, 2023
|publisher=Birdfact
}}
{{cite web
|url=https://indianapublicmedia.org/amomentofscience/how-does-the-windhover-hover.php
|title=How does the windhover hover?
|author=JOSHUA RAWLEIGH
|date=April 4, 2023
|publisher=Indiana Public Media
}}
|last=Caswell
|first=Brandon
|date=Dec 29, 2021
|url=https://www.thegazette.com/sports/hovering-raptors-in-iowa/
|title=Hovering raptors in Iowa
|newspaper=The Gazette
}}
Elanine kites also engage in "windhovering"; this behaviour is also called "kiting" due the common names of this genus.
{{cite web
|url=https://ebird.org/species/bkskit1
|title=Black-winged Kite Elanus caeruleus
|publisher=Cornell Lab of Ornithology
}}
{{cite web
|url=https://ebird.org/species/auskit1
|title=Black-shouldered Kite Elanus axillaris
|publisher=Cornell Lab of Ornithology
}}
{{cite web
|url=https://www.audubon.org/field-guide/bird/northern-harrier
|title=Northern Harrier Circus hudsonius
|publisher=National Audubon Society
}}
Common kestrel hovering.jpg|Common kestrel
Falco cenchroides 2.jpg|Nankeen kestrel
Northern Harrier - 51872832791.jpg|Northern harrier
Black-winged (black-shouldered) kite, Elanus caeruleus, at Marievale, Gauteng, South Africa (45979034625).jpg|Black-winged kite
Letter-winged Kite (Elanus scriptus)4.jpg|Letter-winged kite
White-Tailed Kite.JPG|White-tailed kite
=Seabirds=
Certain seabirds can windhover by soaring or flapping into the wind; often this behaviour takes advantage of thermals whipping off a coastal cliff.
{{cite journal
|url=https://www.vqronline.org/essay/flight-seabirds
|title=The Flight of Seabirds
|author=Louis J. Halle
|date=December 12, 2003
|volume=55
|issue=1
|journal=The Virginia Quarterly Review
}}
{{cite journal
|url=https://birdsoftheworld.org/bow/species/norful/cur/introduction
|title=Northern Fulmar Fulmarus glacialis
|date=March 4, 2020
|last1=Mallory
|first1=Mark L.
|last2=Hatch
|first2=Scott A.
|last3=Nettleship
|first3=David N.
|editor-first1=Shawn M.
|editor-last1=Billerman
|journal=Birds of the World
|doi=10.2173/bow.norful.01
|s2cid=216353101
|url-access=subscription
}}
Tropicbirds can even fly backwards against a strong headwind; Red-tailed tropicbird pairs use this ability to circle each other during courtship displays.
{{cite web
|url=https://www.oiseaux-birds.com/card-red-tailed-tropicbird.html
|title=Red-tailed Tropicbird Phaethon rubricauda
|author=Nicole Bouglouan
|publisher=Oiseaux-Birds
}}
{{cite web
|url=https://ebird.org/species/rettro
|title=Red-tailed Tropicbird Phaethon rubricauda
|publisher=Cornell Lab of Ornithology
}}
Smaller seabirds such as shearwaters and storm petrels feed by hovering low over the water surface,
{{cite web
|url=https://www.allaboutbirds.org/guide/Fork-tailed_Storm-Petrel/overview
|title=Fork-tailed Storm-Petrel Hydrobates furcatus
|publisher=Cornell University
}}
flapping with half-open wings and paddling with their feet in a technique called "pattering" or "sea-anchoring".
|last1=Steele
|first1=Martha
|url=https://sora.unm.edu/node/141755
|title=Meet Our Cover Artist
|date=1992
|journal=Bird Observer
|volume=20
|issue=4
|page=233
|publisher=University of New Mexico
}}
|last1=Xue
|first1=Jiaqi
|last2=Han
|first2=Fei
|last3=van Oorschot
|first3=Brett Klaassen
|last4=Clifton
|first4=Glenna T
|url=https://www.researchgate.net/publication/374487557
|title=Exploring storm petrel pattering and sea-anchoring using deep reinforcement learning
|date=October 2023
|journal=Bioinspiration & Biomimetics
|volume=18
|issue=6
|publisher=University of Portland
|via=ResearchGate
|doi=10.1088/1748-3190/ad00a2
|pmid=37797650
|bibcode=2023BiBi...18f6016X
|s2cid=263705229
|doi-access=free
}}
The waves are accompanied by a slight horizontal wind that enables the birds to soar in place while using their feet to steady themselves.
|url=https://www.researchgate.net/publication/237815118
|journal=Journal of Experimental Biology
|year=1979
|volume=80
|pages=83–91
|title=Aerodynamics and hydrodynamics of the 'hovering' flight of Wilson's Storm Petrel
|last= Withers
|first=Philip
|via=ResearchGate
|doi=10.1242/jeb.80.1.83
}}
Fulmarus glacialis-hovering.jpg|Northern fulmar
Fregata magnificens-female hovering.jpg|Magnificent frigatebird
Phaethon_rubricauda-hovering_flying_backwards.jpg|Red-tailed tropicbird flying backwards
Phaethon rubricauda-courtship_fly_backwards.jpg|Red-tailed tropicbirds circling courtship
Oceanites oceanicus - SE Tasmania.jpg|Wilson's storm petrel pattering
Puffinus gavia-pattering.jpg|Fluttering shearwater pattering