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Flying squirrel
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==Description== [[File:Flying squirrel in a tree.jpg|left|thumb|A [[northern flying squirrel]] (''Glaucomys sabrinus'') gliding]] [[File:Glaucomys volans 1zz.jpg|left|thumb|[[Southern flying squirrel]] (''Glaucomys volans'') skeleton at the [[National Museum of Natural History|NMNH]]. The styliform cartilage attached to the wrist supports the wing membrane.<ref name=":2" />]] Flying squirrels are not capable of flight like [[bird]]s or [[bat]]s; instead, they [[gliding (flight)|glide]] between trees. They are capable of obtaining lift within the course of these flights, with flights recorded to {{convert|90|m|ft}}.<ref name=squirrel>{{cite web |author1=Malamuth, E. |author2=Mulheisen, M. |title=ADW: Glaucomys sabrinus β Northern flying squirrel |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Glaucomys_sabrinus.html |date=1995β2008 |publisher=University of Michigan Museum of Natural History |access-date=14 July 2009 |url-status=live |archive-url=https://web.archive.org/web/20090812040048/http://animaldiversity.ummz.umich.edu/site/accounts/information/Glaucomys_sabrinus.html |archive-date=12 August 2009 }}</ref><ref name=siberian>{{cite journal |last=Asari |first=Y |author2=Yanagawa, H. |author3=Oshida, T. |year=2007 |title=Gliding ability of the Siberian flying squirrel ''Pteromys volans orii'' |journal=Mammal Study |volume=32 |pages=151β154 |url=http://cstl-csm.semo.edu/scheibe/Advanced%20Ecology/Pteromys%20Gliding.pdf |archive-url=https://web.archive.org/web/20100711135035/http://cstl-csm.semo.edu/scheibe/Advanced%20Ecology/Pteromys%20Gliding.pdf |url-status=dead |archive-date=2010-07-11 |access-date=2009-07-14 |doi=10.3106/1348-6160(2007)32[151:GAOTSF]2.0.CO;2 |issue=4 |s2cid=59025388 }}</ref> The direction and speed of the animal in midair are varied by changing the positions of its limbs, largely controlled by small [[cartilaginous]] wrist bones. There is a cartilage projection from the wrist that the squirrel holds upwards during a glide.<ref name=":1" /> This specialized cartilage is only present in flying squirrels and not other gliding mammals.<ref name=":2">{{Cite journal|last1=Kawashima|first1=Tomokazu|last2=Thorington|first2=Richard W.|last3=Bohaska|first3=Paula W.|last4=Sato|first4=Fumi|date=2017-02-01|title=Evolutionary Transformation of the Palmaris Longus Muscle in Flying Squirrels (Pteromyini: Sciuridae): An Anatomical Consideration of the Origin of the Uniquely Specialized Styliform Cartilage|journal=The Anatomical Record|language=en|volume=300|issue=2|pages=340β352|doi=10.1002/ar.23471|pmid=27611816|s2cid=3628991|issn=1932-8494|doi-access=free}}</ref> Possible origins for the styliform cartilage have been explored, and the data suggests that it is most likely homologous to the carpal structures that can be found in other squirrels.<ref name=":2" /> This cartilage along with the manus forms a wing tip to be used during gliding. After being extended, the wing tip may adjust to various angles, controlling aerodynamic movements.<ref>{{Cite journal|title = Myology of the Gliding Membranes of Some Petauristine Rodents (Genera: Glaucomys, Pteromys, Petinomys, and Petaurista)|last = Johnson-Murray|first = Jane L.|date = 1977|journal = Journal of Mammalogy|doi = 10.2307/1379336|volume = 58|issue = 3|pages = 374β384|jstor = 1379336}}</ref><ref name=aerodynamic>{{cite journal |last=Thorington |first=R.W Jr. |author2=Darrow, K. |author3=Anderson, C.G. |year=1998 |title=Wing Tip Anatomy and Aerodynamics in Flying Squirrels |journal=Journal of Mammalogy |volume=79 |issue=1 |pages=245β250 |url=http://entomology.si.edu/StaffPages/Darrow/1997WingTipAnatomy.pdf |access-date=2009-07-14 |doi=10.2307/1382860 |jstor=1382860 |url-status=live |archive-url=https://web.archive.org/web/20090409052036/http://entomology.si.edu/StaffPages/Darrow/1997WingTipAnatomy.pdf |archive-date=2009-04-09 |doi-access=free }}</ref> The wrist also changes the tautness of the [[patagium]], a furry parachute-like membrane that stretches from wrist to ankle.<ref name=aerodynamic/> It has a fluffy tail that stabilizes in flight. The tail acts as an adjunct [[airfoil]], working as an [[Air brake (aircraft)|air brake]] before landing on a tree trunk.<ref>{{cite journal |last=Carraway |first=L.N. |author2=Verts, B.J. |year=1994 |title=Sciurus griseus |journal=Mammalian Species |pages=1β7 |url=http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-474-01-0001.pdf |access-date=2009-07-14 |doi=10.2307/3504097 |issue=474 |url-status=live |archive-url=https://web.archive.org/web/20100107120344/http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-474-01-0001.pdf |archive-date=2010-01-07 |jstor=3504097 |s2cid=253911325 }}</ref> ===Similar gliding animals=== The [[colugo]]s, [[Petauridae]], and [[Anomaluridae]] are gliding mammals which are similar to flying squirrels through [[convergent evolution]], although are not particularly close in relation. Like the flying squirrel, they are [[scansorial]] mammals that use their patagium to glide, unpowered, to move quickly through their environment. ===Evolutionary history=== Prior to the 21st century, the evolutionary history of the flying squirrel was frequently debated.<ref>{{cite journal|last=Arbogast|first=B.S.|title=A brief history of the new world flying squirrels: Phylogeny, biogeography, and conservation genetics|journal=Journal of Mammalogy|year=2007|volume=88|issue=4|pages=840β849|doi=10.1644/06-MAMM-S-322R1.1|doi-access=free}}</ref> This debate was clarified greatly as a result of two molecular studies.<ref>{{cite journal|last=Mercer|first=J.M.|author2=V.L. Roth|s2cid=40366357|title=The effects of cenozoic global change on squirrel phylogeny|journal=Science|year=2003|volume=299|issue=5612|pages=1568β1572|doi=10.1126/science.1079705|pmid=12595609|bibcode=2003Sci...299.1568M}}</ref><ref>{{cite journal|last=Steppan|first=S.J. |author2=B.L. Storz |author3=R.S. Hoffmann|title=Nuclear DNA phylogeny of the squirrels (Mammalia : Rodentia) and the evolution of arboreality from c-myc and RAG1|journal=Molecular Phylogenetics and Evolution|year=2004|volume=30|issue=3|pages=703β719|doi=10.1016/S1055-7903(03)00204-5|pmid=15012949|bibcode=2004MolPE..30..703S }}</ref> These studies found support that flying squirrels originated 18β20 million years ago, are monophyletic, and have a sister relationship with tree squirrels. Due to their close ancestry, the morphological differences between flying squirrels and tree squirrels reveal insight into the formation of the gliding mechanism. Compared to squirrels of similar size, flying squirrels, northern and southern flying squirrels show lengthening in bones of the lumbar vertebrae and forearm, whereas bones of the feet, hands, and distal vertebrae are reduced in length. Such differences in body proportions reveal the flying squirrels' adaptation to minimize wing loading and to increase maneuverability while gliding. The consequence for these differences is that unlike regular squirrels, flying squirrels are not well adapted for quadrupedal locomotion and therefore must rely more heavily on their gliding abilities.<ref>{{Cite journal|title = How to make a flying squirrel: Glaucomys anatomy in phylogenetic perspective|last1 = Thorington|first1 = Richard W.|date = 2007|journal = Journal of Mammalogy|doi = 10.1644/06-mamm-s-325r2.1|first2 = Erica M.|last2 = Santana|volume = 88|issue = 4|pages = 882β896|doi-access = free}}</ref> Several hypotheses have attempted to explain the evolution of gliding in flying squirrels.<ref>{{cite journal|last=Flaherty|first=E.A. |author2=M. Ben-David |author3=W.P. Smith|title=Quadrupedal locomotor performance in two species of arboreal squirrels: predicting energy savings of gliding|journal=Journal of Comparative Physiology B|year=2010|volume=180|issue=7|pages=1067β1078|doi=10.1007/s00360-010-0470-1|pmid=20361193 |s2cid=240833 }}</ref> One possible explanation is related to energy efficiency and foraging.<ref name=":0">{{Cite journal|title = Evolution of vertebrate flight: an aerodynamic model for the transition from gliding to active flight|last = Norberg|first = Ulla M.|date = 1985|journal = American Naturalist|doi = 10.1086/284419|pages = 303β327|volume = 126|issue = 3|s2cid = 85306259}}</ref><ref name=":1">{{Cite journal|title = Take-off and landing forces and the evolution of controlled gliding in northern flying squirrels Glaucomys sabrinus|last1 = Paskins|first1 = Keith E.|date = 2007|journal = The Journal of Experimental Biology|doi = 10.1242/jeb.02747|pmid = 17401124|last2 = Bowyer|first2 = Adrian|last3 = Megill|last4 = Scheibe|first3 = William M.|first4 = John S.|pages = 1413β1423|volume = 210|issue = Pt 8|doi-access = free}}</ref> Gliding is an energetically efficient way to progress from one tree to another while foraging, as opposed to climbing down trees and maneuvering on the ground floor or executing dangerous leaps in the air.<ref name=":0" /> By gliding at high speeds, flying squirrels can rummage through a greater area of forest more quickly than tree squirrels. Flying squirrels can glide long distances by increasing their aerial speed and increasing their lift.<ref name=":1" /> Other hypotheses state that the mechanism evolved to avoid nearby predators and prevent injuries. If a dangerous situation arises on a specific tree, flying squirrels can glide to another, and thereby typically escape the previous danger.<ref name=":1" /><ref>{{Cite journal|title = Morphological attributes of gliding rodents: a preliminary analysis|last1 = Scheibe|first1 = John S.|date = 1990|journal = Transactions of the Missouri Academy of Science|last2 = Figgs|first2 = Daylan|first3 = Jeff|last3 = Heiland|volume = 24|pages = 49β56}}</ref> Furthermore, take-off and landing procedures during leaps, implemented for safety purposes, may explain the gliding mechanism. While leaps at high speeds are important to escape danger, the high-force impact of landing on a new tree could be detrimental to a squirrel's health.<ref name=":1" /> Yet the gliding mechanism of flying squirrels involves structures and techniques during flight that allow for great stability and control. If a leap is miscalculated, a flying squirrel may easily steer back onto the original course by using its gliding ability.<ref name=":1" /> A flying squirrel also creates a large glide angle when approaching its target tree, decreasing its velocity due to an increase in air resistance and allowing all four limbs to absorb the impact of the target.<ref name=":1" /><ref>{{Cite journal|title = Ecological and Biomechanical Insights into the Evolution of Gliding in Mammals|last1 = Byrnes|first1 = Greg|date = 2011|journal = Integrative and Comparative Biology|doi = 10.1093/icb/icr069|pmid = 21719434|last2 = Spence|first2 = Andrew J.|pages = 991β1001|volume = 51|issue = 6|doi-access = free}}</ref> === Fluorescence === In 2019 it was observed, by chance, that a flying squirrel [[Fluorescence|fluoresced]] pink under UV light. Subsequent research by biologists at [[Northland College (Wisconsin)|Northland College]] in Northern [[Wisconsin]] found that this is true for all three species of North American flying squirrels. At this time it is unknown what purpose this serves. Non-flying squirrels do not fluoresce under UV light.<ref>{{Cite journal|last1=Anich|first1=Paula Spaeth|last2=Martin|first2=Jonathan G.|last3=Olson|first3=Erik R.|last4=Kohler|first4=Allison M.|title=Ultraviolet fluorescence discovered in New World flying squirrels (Glaucomys)|journal=Journal of Mammalogy|volume=100|pages=21β30|language=en|doi=10.1093/jmammal/gyy177|year=2019}}</ref>
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