Editing Frog (section)

==Locomotion==
Different species of frog use a number of methods of moving around including [[jumping]], [[running]], [[walking]], [[aquatic locomotion|swimming]], [[burrow]]ing, [[climbing]] and [[Flying frog|gliding]].

[[File:Colostethus flotator jumping.jpg|thumb|left|[[Rainforest rocket frog]] jumping]]
=== Jumping ===
Frogs are generally recognised as exceptional jumpers and, relative to their size, the best jumpers of all vertebrates.<ref>{{cite web |url=http://scienceray.com/biology/zoology/top-10-best-jumper-animals/ |title=Top 10 best jumper animals |publisher=Scienceray |access-date=June 11, 2012 |url-status=dead |archive-url=https://web.archive.org/web/20120810065127/http://scienceray.com/biology/zoology/top-10-best-jumper-animals/ |archive-date=August 10, 2012 }}</ref> The [[striped rocket frog]], ''Litoria nasuta'', can leap over {{convert|2|m|ft|frac=2|abbr=off|spell=in}}, a distance that is more than fifty times its body length of {{convert|55|mm|in|frac=4|abbr=on}}.<ref>{{cite journal |author1=James, R. S. |author2=Wilson, R. S. |year=2008 |title=Explosive jumping: extreme morphological and physiological specializations of Australian rocket frogs (''Litoria nasuta'') |journal=Physiological and Biochemical Zoology |volume=81 |issue=2 |pages=176–185  |pmid=18190283 |doi=10.1086/525290 |s2cid=12643425 |url=http://espace.library.uq.edu.au/view/UQ:175725/UQ175725_OA.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://espace.library.uq.edu.au/view/UQ:175725/UQ175725_OA.pdf |archive-date=October 9, 2022 |url-status=live }}</ref> There are tremendous differences between species in jumping capability. Within a species, jump distance increases with increasing size, but relative jumping distance (body-lengths jumped) decreases. The [[Euphlyctis cyanophlyctis|Indian skipper frog]] (''Euphlyctis cyanophlyctis'') has the ability to leap out of the water from a position floating on the surface.<ref>{{cite journal |author1=Nauwelaerts, S. |author2=Schollier, J. |author3=Aerts, P. |year=2004 |title=A functional analysis of how frogs jump out of water |journal=Biological Journal of the Linnean Society |volume=83 |issue=3 |pages=413–420 |doi=10.1111/j.1095-8312.2004.00403.x |doi-access=free }}</ref> The tiny [[northern cricket frog]] (''Acris crepitans'') can "skitter" across the surface of a pond with a series of short rapid jumps.<ref name=Badger2>{{cite book |title=Frogs |last=Badger |first=David |author2=Netherton, John |year=1995 |publisher=Airlife Publishing |isbn=978-1-85310-740-5 |page=51 }}</ref>

Slow-motion photography shows that the muscles have passive flexibility. They are first stretched while the frog is still in the crouched position, then they are contracted before being stretched again to launch the frog into the air. The fore legs are folded against the chest and the hind legs remain in the extended, streamlined position for the duration of the jump.<ref name=Frogjump>{{cite magazine |url=http://www.komverse.com/2010/05/15/how-frogs-jump-nationalgeographic/ |title=How frogs jump |author=Minott, Kevin |date=May 15, 2010 |magazine=National Geographic |access-date=June 10, 2012 |archive-url=https://web.archive.org/web/20131104045512/http://www.komverse.com/2010/05/15/how-frogs-jump-nationalgeographic/ |archive-date=November 4, 2013 |url-status=dead }}</ref> In some extremely capable jumpers, such as the [[Cuban tree frog]] (''Osteopilus septentrionalis'') and the [[northern leopard frog]] (''Lithobates pipiens''), the peak power exerted during a jump can exceed that which the muscle is theoretically capable of producing. When the muscles contract, the energy is first transferred into the stretched tendon which is wrapped around the ankle bone. Then the muscles stretch again at the same time as the tendon releases its energy like a [[catapult]] to produce a powerful acceleration beyond the limits of muscle-powered acceleration.<ref>{{cite journal |author1=Astley, H. C. |author2=Roberts, T. J. |year=2011 |title=Evidence for a vertebrate catapult: elastic energy storage in the plantaris tendon during frog jumping |journal=Biology Letters |volume=8 |issue=3 |pages=386–389 |doi=10.1098/rsbl.2011.0982 |pmid=22090204 |pmc=3367733 }}</ref> A similar mechanism has been documented in [[locust]]s and [[grasshopper]]s.<ref>{{cite journal |last1=Scott |first1=J. |title=The locust jump: an integrated laboratory investigation |doi=10.1152/advan.00037.2004 |year=2005 |pages=21–26 |volume=29 |journal=Advances in Physiology Education |issue=1 |pmid=15718379|s2cid=13423666 }}</ref>

Early hatching of froglets can have negative effects on frog jumping performance and overall locomotion.<ref name=":1">{{Cite journal|last1=Buckley|first1=C. R.|last2=Michael|first2=S. F.|last3=Irschick|first3=D. J.|date=2005|title=Early Hatching Decreases Jumping Performance in a Direct-Developing Frog, Eleutherodactylus coqui|journal=Functional Ecology|volume=19|issue=1|pages=67–72|doi=10.1111/j.0269-8463.2005.00931.x|jstor=3599272|issn=0269-8463|doi-access=free|bibcode=2005FuEco..19...67B }}</ref> The hindlimbs are unable to completely form, which results in them being shorter and much weaker relative to a normal hatching froglet.<ref name=":1" /> Early hatching froglets may tend to depend on other forms of locomotion more often, such as swimming and walking.<ref name=":1" />

=== Walking and running ===
[[File:Phrynosequence bodyFoR2.gif|thumb|right|''[[Phrynomantis bifasciatus]]'' walking on a level surface]]
Frogs in the families Bufonidae, [[Rhinophrynidae]], and [[Microhylidae]] have short back legs and tend to walk rather than jump.<ref name=Britannica>{{cite encyclopedia |last1=Zug |first1=George R. |last2=Duellman |first2=William E. |encyclopedia=Encyclopædia Britannica Online |title=Anura |url=https://www.britannica.com/EBchecked/topic/29023/Anura |access-date=April 26, 2015 |date=May 14, 2014 }}</ref> When they try to move rapidly, they speed up the rate of movement of their limbs or resort to an ungainly hopping gait. The [[Great Plains narrow-mouthed toad]] (''Gastrophryne olivacea'') has been described as having a gait that is "a combination of running and short hops that are usually only an inch or two in length".<ref>{{cite journal |author=Fitch, H. S. |year=1956 |title=An ecological study of the collared lizard (''Crotaphytus collaris'') |journal=University of Kansas Publications  |volume=8 |pages=213–274 }}</ref> In an experiment, [[Bufo fowleri|Fowler's toad]] (''Anaxyrus'' ''fowleri'') was placed on a treadmill which was turned at varying speeds. By measuring the toad's uptake of oxygen it was found that hopping was an inefficient use of resources during sustained locomotion but was a useful strategy during short bursts of high-intensity activity.<ref>{{cite journal |author1=Walton, M. |author2=Anderson, B. D. |year=1988 |title=The aerobic cost of saltatory locomotion in the fowler's toad (''Bufo woodhousei fowleri'') |journal=Journal of Experimental Biology |volume=136 |pages=273–288 |url=http://jeb.biologists.org/content/136/1/273 |pmid= 3404074 |issue=1|doi=10.1242/jeb.136.1.273 |doi-access=free |bibcode=1988JExpB.136..273W }} {{free access}}</ref>

The [[Kassina maculata|red-legged running frog]] (''Kassina maculata'') has short, slim hind limbs unsuited to jumping. It can move fast by using a running gait in which the two hind legs are used alternately. Slow-motion photography shows, unlike a horse that can trot or gallop, the frog's gait remained similar at slow, medium, and fast speeds.<ref>{{cite journal |author1=Ahn, A. N. |author2=Furrow, E. |author3=Biewener, A. A. |year=2004 |title=Walking and running in the red-legged running frog, ''Kassina maculata'' |journal=Journal of Experimental Biology |volume=207 |pages=399–410 |doi=10.1242/jeb.00761 |pmid= 14691087 |issue= Pt 3 |doi-access=free |bibcode=2004JExpB.207..399A }}</ref> This species can also climb trees and shrubs, and does so at night to catch insects.<ref>{{cite iucn |url=https://www.iucnredlist.org/species/56231/11445543 |title=''Kassina maculata'' |author1=Pickersgill, M. |author2=Schiøtz, A. |author3=Howell, K. |author4=Minter, L. |year=2004 |access-date=June 11, 2012}}</ref> The Indian skipper frog (''Euphlyctis cyanophlyctis'') has broad feet and can run across the surface of the water for several metres (yards).<ref name=Badger2/>

=== Swimming ===
[[File:Frog žába.gif|thumb|alt=Common toad swimming|[[Common toad]] (''Bufo bufo'') swimming]]

Frogs that live in or visit water have adaptations that improve their swimming abilities. The hind limbs are heavily muscled and strong. The webbing between the toes of the hind feet increases the area of the foot and helps propel the frog powerfully through the water. Members of the family [[Pipidae]] are wholly aquatic and show the most marked specialisation. They have inflexible vertebral columns, flattened, streamlined bodies, [[lateral line]] systems, and powerful hind limbs with large webbed feet.<ref name=Pipidae>{{cite web |url=http://amphibiaweb.org/lists/Pipidae.shtml |title=Pipidae |work=AmphibiaWeb |publisher=University of California, Berkeley |access-date=June 14, 2012}}</ref> Tadpoles mostly have large tail fins which provide thrust when the tail is moved from side to side.<ref name="Duellman"/>

=== Burrowing ===
Some frogs have become adapted for burrowing and a life underground. They tend to have rounded bodies, short limbs, small heads with bulging eyes, and hind feet adapted for excavation. An extreme example of this is the [[purple frog]] (''Nasikabatrachus sahyadrensis'') from southern India which feeds on [[termite]]s and spends almost its whole life underground. It emerges briefly during the [[monsoon]] to mate and breed in temporary pools. It has a tiny head with a pointed snout and a plump, rounded body. Because of this [[fossorial]] existence, it was [[Species description|first described]] in 2003, being new to the scientific community at that time, although previously known to local people.<ref>{{cite journal |author1=Radhakrishnan, C. |author2=Gopi, K. C. |year=2007 |title=Extension of range of distribution of ''Nasikabatrachus sahyadrensis'' Biju & Bossuyt ( Amphibia : Anura : Nasikabatrachidae ) along Western Ghats, with some insights into its bionomics |journal=Current Science |volume=92 |issue=2 |pages=213–216 |issn=0011-3891 |url=http://www.iisc.ernet.in/currsci/jan252007/213.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.iisc.ernet.in/currsci/jan252007/213.pdf |archive-date=October 9, 2022 |url-status=live }}</ref>

[[File:Nasikabatrachus sahyadrensis.jpg|thumb|left|alt=Purple frog|[[Purple frog]] (''Nasikabatrachus sahyadrensis'')]]

The spadefoot toads of North America are also adapted to underground life. The [[Plains spadefoot toad]] (''Spea bombifrons'') is typical and has a flap of keratinised bone attached to one of the [[Metatarsus|metatarsals]] of the hind feet which it uses to dig itself backwards into the ground. As it digs, the toad wriggles its hips from side to side to sink into the loose soil. It has a shallow burrow in the summer from which it emerges at night to forage. In winter, it digs much deeper and has been recorded at a depth of {{convert|4.5|m|ftin|abbr=on}}.<ref name=Farrar>{{cite web |url=http://amphibiaweb.org/cgi-bin/amphib_query?where-genus=Spea&where-species=bombifrons |title=''Spea bombifrons''  |author1=Farrar, Eugenia |author2=Hey, Jane |work=AmphibiaWeb |publisher=University of California, Berkeley |access-date=June 16, 2012}}</ref>  The tunnel is filled with soil and the toad hibernates in a small chamber at the end. During this time, urea accumulates in its tissues and water is drawn in from the surrounding damp soil by [[osmosis]] to supply the toad's needs.<ref name=Farrar/> Spadefoot toads are explosive breeders, all emerging from their burrows at the same time and converging on temporary pools, attracted to one of these by the calling of the first male to find a suitable breeding location.<ref>{{cite web |url=http://amphibiaweb.org/lists/Scaphiopodidae.shtml |title=Scaphiopodidae |work=AmphibiaWeb |publisher=University of California, Berkeley |access-date=June 16, 2012}}</ref>

The burrowing frogs of Australia have a different lifestyle. The [[western spotted frog]] (''Heleioporus albopunctatus'') digs a burrow beside a river or in the bed of an ephemeral stream and regularly emerges to forage. Mating takes place and eggs are laid in a foam nest inside the burrow. The eggs partially develop there but do not hatch until they are submerged following heavy rainfall. The tadpoles then swim out into the open water and rapidly complete their development.<ref>{{cite iucn |url=https://www.iucnredlist.org/species/9763/13014492 |title=''Heleioporus albopunctatus'' |author1=Roberts, Dale |author2=Hero, Jean-Marc |year=2011 |access-date=June 16, 2012}}</ref> Madagascan burrowing frogs are less fossorial and mostly bury themselves in leaf litter. One of these, the [[Scaphiophryne marmorata|green burrowing frog]] (''Scaphiophryne marmorata''), has a flattened head with a short snout and well-developed metatarsal tubercles on its hind feet to help with excavation. It also has greatly enlarged terminal discs on its fore feet that help it to clamber around in bushes.<ref>{{cite web |url=http://www.amphibian.co.uk/scaphiophryne.html |title=Madagascan Burrowing Frogs: Genus: ''Scaphiophryne'' (Boulenger, 1882) |author=Staniszewski, Marc |date=September 30, 1998 |access-date=June 16, 2012}}</ref>  It breeds in temporary pools that form after rains.<ref>{{cite journal |author1=Venesci, M |author2=Raxworthy, C. J. |author3=Nussbaum, R. A. |author4=Glaw, F. |year=2003 |title=A revision of the ''Scaphiophryne marmorata'' complex of marbled toads from Madagascar, including the description of a new species |journal=Herpetological Journal |volume=13 |pages=69–79 |url=http://www.mvences.de/p/p1/Vences_A51.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.mvences.de/p/p1/Vences_A51.pdf |archive-date=October 9, 2022 |url-status=live }}</ref>

=== Climbing ===
[[File:Phyllomedusa burmeisteri01.jpg|thumb|alt=Frog climbing|[[Phyllomedusa burmeisteri|Burmeister's leaf frog]]]]
[[File:Glass frogs.jpg|thumb|Group of [[glass frog]]s]]
Tree frogs live high in the [[canopy (biology)|canopy]], where they scramble around on the branches, twigs, and leaves, sometimes never coming down to earth. The "true" tree frogs belong to the family Hylidae, but members of other frog families have independently adopted an arboreal habit, a case of [[convergent evolution]]. These include the [[glass frogs]] (Centrolenidae), the [[Hyperoliidae|bush frogs]] (Hyperoliidae), some of the narrow-mouthed frogs (Microhylidae), and the [[Rhacophoridae|shrub frogs]] (Rhacophoridae).<ref name=Britannica/> Most tree frogs are under {{convert|10|cm|in|0|abbr=on}} in length, with long legs and long toes with adhesive pads on the tips. The surface of the toe pads is formed from a closely packed layer of flat-topped, hexagonal [[epidermis (skin)|epidermal]] cells separated by grooves into which glands secrete [[mucus]]. These toe pads, moistened by the mucus, provide the grip on any wet or dry surface, including glass. The forces involved include [[boundary friction]] of the toe pad epidermis on the surface and also [[surface tension]] and [[viscosity]].<ref>{{cite journal |author1=Federle, W. |author2=Barnes, W. J. P. |author3=Baumgartner, W. |author4=Drechsler, P. |author5=Smith, J. M. |year=2006 |title=Wet but not slippery: boundary friction in tree frog adhesive toe pads |journal=Journal of the Royal Society Interface |volume=3 |issue=10 |pages=689–697 |doi=10.1098/rsif.2006.0135 |pmc=1664653 |pmid=16971337}}</ref> Tree frogs are very acrobatic and can catch insects while hanging by one toe from a twig or clutching onto the blade of a windswept reed.<ref>{{cite book |title=Living Amphibians of the World |last=Cochran |first=Doris Mabel |year=1961 |publisher=Doubleday |isbn=978-0-241-90338-4 |page=112 |url={{Google books|HOk6AAAAMAAJ|page=112|plainurl=yes}} }}</ref> Some members of the subfamily [[Phyllomedusinae]] have [[Opposable thumb#Opposition and apposition|opposable toes]] on their feet. The [[Phyllomedusa ayeaye|reticulated leaf frog]] (''Phyllomedusa ayeaye'') has a single opposed [[Digit (anatomy)|digit]] on each fore foot and two opposed digits on its hind feet. This allows it to grasp the stems of bushes as it clambers around in its riverside habitat.<ref>{{cite web |url=http://amphibiaweb.org/cgi-bin/amphib_query?where-genus=Phyllomedusa&where-species=ayeaye |title=''Phyllomedusa ayeaye'' |publisher=AmphibiaWeb |access-date=June 14, 2012}}</ref>

=== Gliding ===
During the evolutionary history of frogs, several different groups have independently taken to the air.<ref>{{cite journal |author1=Emerson, Sharon B. |author2=Koehl, M. A. R.  |year=1990 |title=The interaction of behavioral and morphological change in the evolution of a novel locomotor type: "flying frogs |journal=Evolution |volume=44 |issue=8 |pages=1931–1946 |jstor=2409604 |doi=10.2307/2409604 |pmid=28564439 }}</ref> Some frogs in the tropical rainforest are specially adapted for gliding from tree to tree or parachuting to the forest floor. Typical of them is [[Rhacophorus nigropalmatus|Wallace's flying frog]] (''Rhacophorus nigropalmatus'') from Malaysia and Borneo. It has large feet with the fingertips expanded into flat adhesive discs and the digits fully webbed. Flaps of skin occur on the lateral margins of the limbs and across the tail region. With the digits splayed, the limbs outstretched, and these flaps spread, it can glide considerable distances, but is unable to undertake powered flight.<ref>{{cite web |url=http://amphibiaweb.org/cgi-bin/amphib_query?where-genus=Rhacophorus&where-species=nigropalmatus |title=''Rhacophorus nigropalmatus'' |date=November 29, 2001 |author1=Shah, Sunny |author2=Tiwari, Rachna |work=AmphibiaWeb |publisher=University of California, Berkeley |access-date=June 11, 2012}}</ref> It can alter its direction of travel and navigate distances of up to {{convert|15|m|ft|round=5|abbr=on}} between trees.<ref>{{cite web |url=http://animals.nationalgeographic.com/animals/amphibians/wallaces-flying-frog/ |archive-url=https://web.archive.org/web/20100118144053/http://animals.nationalgeographic.com/animals/amphibians/wallaces-flying-frog |url-status=dead |archive-date=January 18, 2010 |title=Wallace's Flying Frog ''Rhacophorus nigropalmatus'' |publisher=National Geographic: Animals |access-date=June 5, 2012|date=September 10, 2010 }}</ref>