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Hover (behaviour)

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Squash vine borer 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". [1] [2]

True hoverers

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Hummingbirds

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Hummingbirds hover over flowers to obtain nectar, flapping their wings at up to 70 beats per second. [3]

Bats

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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. [4] [5]

Kingfishers

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Small Kingfishers such as Belted kingfisher may hover over water before diving in to catch fish. [6] Larger species such as Ringed kingfisher are too heavy to hover for more than a few seconds. [7]

Moths

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Sphinx moths

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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. [8]

Clearwing moths

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Some clearwing moths (family Sesiidae) also hover while nectaring [9] [10] or even puddling. [11] Females may also hover to inspect ovipositing sites. [12]

Hoverflies

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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,[13] while female hovering serves to inspect ovipositing sites. [14] [15] [16]

Bee flies

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Bee flies are parasitoids that can dart about in the air with great agility. Males hover as a courtship display, [17] [18] 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. [19] 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. [20]

Odonata

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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. [21]

Dragonflies

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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. [22]

Damselflies

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Some male damselflies hover in front of females or over the oviposition site during courtship; sometimes females also hover in response. [23] [24] 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. [25] [26]

Hymenoptera

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Bees

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Many bee species, such as bumblebees, hover momentarily as they approach flowers to feed. [27] Males of some species, including carpenter bees and carder bees, also hover while patrolling their territories. [28] [29] [30]

Wasps

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Among the social wasps, Stenogastrinae are known as hover wasps due to their distinctive hovering flight. [31] Males often hover to display banding patterns on their abdomen as a territorial display. [32] [33]

Among the solitary wasps, parasitoid species such as scoliid wasps exhibit hovering behaviour while hunting for prey to feed their larvae. [34] [35] Males of some parasitoids may hover briefly while they patrol their territories, seeking females and chasing away rivals. [36] [37] [38]

Wind hoverers

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Raptors

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Many birds of prey such as kestrels, harriers, and members of the Buteo genus can "windhover" by facing the wind. [39] [40] [41] Elanine kites also engage in "windhovering"; this behaviour is also called "kiting" due the common names of this genus. [42] [43] [44]

Seabirds

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Certain seabirds can windhover by soaring or flapping into the wind; often this behaviour takes advantage of thermals whipping off a coastal cliff. [45] [46]

Tropicbirds can even fly backwards against a strong headwind; Red-tailed tropicbird pairs use this ability to circle each other during courtship displays. [47] [48]

Smaller seabirds such as shearwaters and storm petrels feed by hovering low over the water surface, [49] flapping with half-open wings and paddling with their feet in a technique called "pattering" or "sea-anchoring". [50] [51] The waves are accompanied by a slight horizontal wind that enables the birds to soar in place while using their feet to steady themselves. [52]

References

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  1. ^ Ehrlich, Paul R.; Dobkin, David S.; Wheye, Darryl. "Hovering Flight". stanfordbirds.
  2. ^ Lekwa, Steve (Feb 22, 2021). "Kiting is a common activity for Iowa's raptors". Ames Tribune.
  3. ^ "The Hummingbird Wing Beat Challenge". National Audubon Society. April 22, 2020.
  4. ^ "Stanford engineers study hovering bats and hummingbirds in Costa Rica". Stanford news. September 26, 2018.
  5. ^ Susan Milius (October 15, 2018). "How nectar bats fly nowhere". Science News.
  6. ^ "Belted Kingfisher Megaceryle alcyon". National Audubon Society.
  7. ^ "Ringed Kingfisher Megaceryle torquata". Cornell University.
  8. ^ "Hummingbird Moth (Clearwing Moth)". Mass Audubon.
  9. ^ BROWN, LARRY N. and; MIZELL, III, RUSSELL F. (1993). "THE CLEARWING BORERS OF FLORIDA (LEPIDOPTERA: SESIIDAE)". Tropical Lepidoptera. 4 (3). Florida Online Journals: 1–21.
  10. ^ Potter, Daniel (2 October 2014). Capinera, J.L. (ed.). "Clearwing Moths (Lepidoptera: Sesiidae)". Encyclopedia of Entomology. Springer, Dordrecht: 928. doi:10.1007/978-1-4020-6359-6_5123. ISBN 978-1-4020-6242-1.
  11. ^ "Melittia celebica". iNaturalist. 4 July 2023.
  12. ^ "Pyropteron icteropus". iNaturalist. 22 May 2022.
  13. ^ Collett, T.S.; Land, M.F. (September 1978). "How hoverflies compute interception courses". Journal of Comparative Physiology. 125 (3). Springer-Verlag: 191–204. doi:10.1007/BF00656597. S2CID 26039329.
  14. ^ Almohamad, Raki; Verheggen, François J.; HaubrugeUniv, Éric (2009). "Searching and oviposition behavior of aphidophagous hoverflies (Diptera: Syrphidae): a review". Biotechnologie, Agronomie, Société et Environnement. 13 (3): 467–481.
  15. ^ Vera Strader. "Hover Flies, a Gardener's Friend" (PDF). University of California Agriculture and Natural Resources.
  16. ^ Peter Chen (December 6, 2023). "Allograpta exotica ovipositing - Allograpta exotica - Female". Bugguide. Iowa State University.
  17. ^ "Bee-flies Bombylius sp" (PDF). Natural History Museum.
  18. ^ Ferguson, David J; Yeates, David K (May 2013). "The courtship behavior of the bee fly Meomyia vetusta Walker (Diptera: Bombyliidae)". Australian Entomologist. 40: 89–92 – via ResearchGate.
  19. ^ Boesi, Roberto; Polidori, Carlo; Andrietti, Francesco (March 2009). "Searching for the Right Target: Oviposition and Feeding Behavior in Bombylius Bee Flies (Diptera: Bombyliidae)". Zoological Studies. 48 (2): 141–150 – via ResearchGate.
  20. ^ "The Large Bee Fly - Bombylius major". Buzzaboutbees. April 20, 2021.
  21. ^ David Britton (Mar 7, 2023). "Dragonflies and damselflies - Order Odonata". Australian Museum.
  22. ^ McMillan, Victoria E. (June 1991). "Variable mate-guarding behaviour in the dragonfly Plathemis lydia (Odonata: Libellulidae)". Animal Behaviour. 41 (6). Elsevier B.V.: 979–987. doi:10.1016/S0003-3472(05)80636-0. S2CID 54360417.
  23. ^ The BugLady (December 14, 2022). "River Jewelwing Damselfly". University of Wisconsin-Milwaukee.
  24. ^ Guillermo-Ferreiraa, Rhainer; Neissc, Ulisses Gaspar; Hamadad, Neusa; Bispo, Pitágoras C. (2 October 2014). "Behavior of the Amazonian damselfly Chalcopteryx scintillans McLachlan (Zygoptera: Polythoridae) and comments on its morphological distinction from C. rutilans (Rambur)". International Journal of Odonatology. 17 (4). Worldwide Dragonfly Association: 251–258. Bibcode:2014IJOdo..17..251G. doi:10.1080/13887890.2014.983189.
  25. ^ Marla Garrison (2011). "Damselflies of Chicagoland A Photo Field Guide, version 2" (PDF). The Field Museum, Chicago.
  26. ^ "Nature Note: Dragonflies and Damselflies". Lake Champlain Committee. August 31, 2009.
  27. ^ "Do Bumble Bees Hover?". Buzzaboutbees.
  28. ^ "Why Do Carpenter Bees Hover?". Best Bee Brothers. January 12, 2023.
  29. ^ MICHAEL F. POTTER. "Carpenter Bees". University of Kentucky.
  30. ^ Samantha Gallagher; Andrea Lucky (December 2019). "common name: European wool carder bee". University of Florida.
  31. ^ Carpenter, James Michael; Nguyen, Lien Thi Phuong (September 2003). "Keys to the genera of social wasps of South-East Asia (Hymenoptera: Vespidae)". Entomological Science. 6 (3). American Museum of Natural History: 183–192. doi:10.1046/j.1343-8786.2003.00016.x. S2CID 86823797 – via ResearchGate.
  32. ^ BEANI, L.; TURILLAZZI, S. (June 1999). "Stripes display in hover-wasps (Vespidae: Stenogastrinae): a socially costly status badge". Animal Behaviour. 57 (6). The Association for the Study of Animal Behaviour: 1233–1239. doi:10.1006/anbe.1999.1101. PMID 10373256. S2CID 45496129 – via Elsevier.
  33. ^ Turillazzi, S.; Francescato, E. (1990). "Patrolling behaviour and related secretory structures in the males of some Stenogastrine wasps". Insectes Sociaux. 37: 146–157. doi:10.1007/BF02224027. S2CID 25746201 – via Springer Nature.
  34. ^ "FLYING PESTS ARE MORE THAN JUST ANNOYING". Daily Press. July 30, 2019.
  35. ^ "Scoliid Wasps - Lawns". University of Maryland Extension. March 1, 2023.
  36. ^ "Eastern Cicada-Killer Wasp". Missouri Department of Conservation.
  37. ^ "Cicada Killer Wasp". N.C. Cooperative Extension.
  38. ^ Connie Schmotzer (July 13, 2015). "Masterful Gardening: Sand wasps - our allies in pest control". YorkDailyRecord.
  39. ^ "Kestrel Hovering: A Complete Guide". Birdfact. March 15, 2023.
  40. ^ JOSHUA RAWLEIGH (April 4, 2023). "How does the windhover hover?". Indiana Public Media.
  41. ^ Caswell, Brandon (Dec 29, 2021). "Hovering raptors in Iowa". The Gazette.
  42. ^ "Black-winged Kite Elanus caeruleus". Cornell Lab of Ornithology.
  43. ^ "Black-shouldered Kite Elanus axillaris". Cornell Lab of Ornithology.
  44. ^ "Northern Harrier Circus hudsonius". National Audubon Society.
  45. ^ Louis J. Halle (December 12, 2003). "The Flight of Seabirds". The Virginia Quarterly Review. 55 (1).
  46. ^ Mallory, Mark L.; Hatch, Scott A.; Nettleship, David N. (March 4, 2020). Billerman, Shawn M. (ed.). "Northern Fulmar Fulmarus glacialis". Birds of the World. doi:10.2173/bow.norful.01. S2CID 216353101.
  47. ^ Nicole Bouglouan. "Red-tailed Tropicbird Phaethon rubricauda". Oiseaux-Birds.
  48. ^ "Red-tailed Tropicbird Phaethon rubricauda". Cornell Lab of Ornithology.
  49. ^ "Fork-tailed Storm-Petrel Hydrobates furcatus". Cornell University.
  50. ^ Steele, Martha (1992). "Meet Our Cover Artist". Bird Observer. 20 (4). University of New Mexico: 233.
  51. ^ Xue, Jiaqi; Han, Fei; van Oorschot, Brett Klaassen; Clifton, Glenna T (October 2023). "Exploring storm petrel pattering and sea-anchoring using deep reinforcement learning". Bioinspiration & Biomimetics. 18 (6). University of Portland. Bibcode:2023BiBi...18f6016X. doi:10.1088/1748-3190/ad00a2. PMID 37797650. S2CID 263705229 – via ResearchGate.
  52. ^ Withers, Philip (1979). "Aerodynamics and hydrodynamics of the 'hovering' flight of Wilson's Storm Petrel". Journal of Experimental Biology. 80: 83–91. doi:10.1242/jeb.80.1.83 – via ResearchGate.
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