A bee in flight.

  • Insects (flight). The first of all animals to evolve flight, insects are also the only invertebrates that have evolved flight. The species are too numerous to list here. Insect flightis an active research field.
    • Gliding bristletails (gliding). Directed aerial gliding descent is found in some tropical arboreal bristletails, an ancestrally wingless sister taxa to the winged insects. The bristletails median caudal filament is important for the glide ratio and gliding control [4]
    • Gliding ants (gliding). The flightless workers of these insects have secondarily gained some capacity to move through the air. Gliding has evolved independently in a number of arboreal ant species from the groups Cephalotini, Pseudomyrmecinae, and Formicinae (mostly Camponotus). All arboreal dolichoderines and non-cephalotine myrmicines except Daceton armigerum do not glide. Living in the rainforest canopy like many other gliders, gliding ants use their gliding to return to the trunk of the tree they live on should they fall or be knocked off a branch. Gliding was first discovered for Cephalotes atreus in the Peruvian rainforest. Cephalotes atreus can make 180 degree turns, and locate the trunk using visual cues, succeeding in landing 80% of the time.[5] Unique among gliding animals, Cephalotini and Pseudomyrmecinae ants glide abdomen first, the Forminicae however glide in the more conventional head first manner.[6] The following page has some good videos of gliding ants. [1]
  • Spiders (parachuting). The young of some species of spiders travel through the air by using silk draglines to catch the wind, as may some smaller species of adult spider, such the money spider family. This behavior is commonly known as “ballooning”. Ballooning spiders make up part of the aeroplankton.

[edit] Molluscs

  • Flying squid (gliding). Several oceanic squids, such as the Pacific flying squid, will leap out of the water to escape predators, an adaptation similar to that of flying fish.[7] Smaller squids will fly in shoals, and have been observed to cover distances as long as 50 meters. Small fins towards the back of the mantle do not produce much lift, but do help stabilize the motion of flight. They exit the water by expelling water out of their funnel, indeed some squid have been observed to continue jetting water while airborne possibly providing thrust even after leaving the water. This may make flying squid the only animals with jet-propelled aerial locomotion.[8]

[edit] Vertebrates

[edit] Fish

Band-winged flying fish. Note the enlarged pectoral fins.

  • Flying fish (gliding). There are over 50 species of flying fish belonging to the family Exocoetidae. They are mostly marine fishes of small to medium size. The largest flying fish can reach lengths of 45 cm, but most species measure less than 30 cm in length. They can be divided into two-winged varieties and four-winged varieties. Before the fish leaves the water it increases its speed to around 30 body lengths per second and as it breaks the surface and is freed from the drag of the water it can be traveling at around 60 km/h.[9] The glides are usually up to 30–50 metres in length, but some have been observed soaring for hundreds of metres using the updraft on the leading edges of waves. The fish can also make a series of glides, each time dipping the tail into the water to produce forward thrust. The longest recorded series of glides, with the fish only periodically dipping its tail in the water, was for 45 seconds (Video here [2]). It has been suggested that the genus Exocoetus is on an evolutionary borderline between flight and gliding. It flaps its enlarged pectoral fins when airborne, but still seems only to glide, as there is no hint of a power stroke.[10] It has been found that some flying fish can glide as effectively as some flying birds [3].
  • Halfbeaks (gliding). A group related to the Exocoetidae, one or two hemirhamphid species possess enlarged pectoral fins and show true gliding flight rather than simple leaps. Marshall (1965) reports that Euleptorhamphus viridis can cover 50 m in two separate hops.[11]
  • Freshwater butterflyfish (possibly gliding). Pantodon buchholzi has the ability to jump and possibly glide a short distance. It can move through the air several times the length of its body. While it does this, the fish flaps its large pectoral fins, giving it its common name.[12] However, it is debated whether the freshwater butterfly fish can truly glide, Saidel et al. (2004) argue that it cannot.
  • Freshwater hatchetfish (possibly flying). There are 9 species of freshwater hatchetfish split among 3 genera. Freshwater hatchetfish have an extremely large sternal region that is fitted with a large amount of muscle that allows it to flap its pectoral fins. They can move in a straight line over a few metres to escape predators.

[edit] Amphibians

  • Rhacophoridae flying frogs (gliding). Gliding has evolved independently in two families of tree frogs, the Old World Rhacophoridae and the New World Hylidae. Within each lineage there are a range of gliding abilities from non-gliding, to parachuting, to full gliding. A number of the Rhacophoridae, such as Wallace’s Flying Frog (Rhacophorus nigropalmatus), have adaptation for gliding, the main feature being enlarged toe membranes. For example, the Malayan flying frog glides using the membranes between the toes of its limbs, and small membranes located at the heel, the base of the leg, and the forearm. Some of the frogs are quite accomplished gliders, for example, the Chinese gliding frog Polypedates dennysi can maneuver in the air, making two kinds of turn, either rolling into the turn (a banked turn) or yawing into the turn (a crabbed turn).
  • Hylidae flying frogs (gliding). The other frog family that contains gliders.

The underside of Kuhl’s flying gecko Ptychozoon kuhli. Note the gliding adaptations: flaps of skin on the legs, feet, sides of the body, and on the sides of the head.

[edit] Reptiles

  • Draco lizards (gliding). There are 28 species of lizard of the genus Draco, found in Sri Lanka, India, and Southeast Asia. They live in trees, feeding on tree ants, but nest on the forest floor. They can glide for up to 60 m and over this distance they lose only 10 m in height.[9] Unusually, their patagium (gliding membrane) is supported on elongated ribs rather than the more common situation among gliding vertebrates of having the patagium attached to the limbs. When extended, the ribs form a semicircle on either side the lizard’s body and can be folded to the body like a folding fan.
  • Gliding Lacertids (gliding). There are two species of gliding lacertid, of the genus Holaspis. Found in Africa. They have fringed toes and tail sides and can flatten their bodies for gliding/parachuting.[13]
  • Ptychozoon flying geckos (gliding). There are six species of gliding gecko, of the genus Ptychozoon, from Southeast Asia. These lizards have small flaps of skin along their limbs, torso, tail, and head that catch the air and enable them to glide. [4]
  • Lupersaurus flying geckos (gliding). A possible sister-taxon to Ptychozoon which has similar flaps and folds and also glides.[5]
  • Thecadactylus flying geckos (gliding). At least some species of Thecadactylus, such as T. rapicauda, are known to glide. [6]
  • Cosymbotus flying gecko (gliding). Similar adaptations to Ptychozoon are found in the two species of the gecko genus Cosymbotus.
  • Chrysopelea snakes (gliding/parachuting). Five species of snake from Southeast Asia, Melanesia, and India. The paradise tree snake of southern Thailand, Malaysia, Borneo, Philippines, and Sulawesi is the most capable glider of those snakes studied. It glides by stretching out its body sideways and opening its ribs so the belly is concave, and by making lateral slithering movements. It can remarkably glide up to 100 m and make 90 degree turns. Follow this link for videos of gliding snakes.

[edit] Birds

Main article: Bird flight

Birds are the most successful group of flying vertebrate.

  • Birds (flying, soaring) Again the species are too numerous to nominate. Bird flight is one of the most studied forms of aerial locomotion in animals. See List of soaring birds for birds that can soar as well as fly.

[edit] Mammals

Townsends’s Big-eared Bat, (Corynorhinus townsendii) displaying the “hand wing”

  • Bats (flying). There are many species of bat, again too numerous to nominate.
  • Flying squirrels (subfamily Petauristinae) (gliding). There are 43 species divided between 14 genera of flying squirrel. Flying squirrels are found almost worldwide in tropical (Southeast Asia, India, and Sri Lanka), temperate, and even Arctic environments. They tend to be nocturnal. When a flying squirrel wishes to cross to a tree that is further away than the distance possible by jumping, it extends the cartilage spur on its elbow or wrist. This opens out the flap of furry skin (the patagium) that stretches from its wrist to its ankle. It glides spread-eagle and with its tail fluffed out like a parachute, and grips the tree with its claws when it lands. Flying squirrels have been reported to glide over 200 m.
  • Anomalure or scaly-tailed flying squirrels (Anomaluridae family) (gliding). These brightly coloured African rodents are not squirrels but have evolved to a resemble flying squirrels by convergent evolution. There are seven species, divided in three genera. All but one species has gliding membranes between their front and hind legs. One genus is particularly small and is known as flying mice, but similarly they are not mice.
  • Colugos or Flying lemurs (order Dermoptera) (gliding). There are two species of flying lemur. This is not a lemur, which is a primate, but molecular evidence suggests that colugos are a sister group to primates, however some mammologists suggest they are a sister group to bats. Found in Southeast Asia, the colugo is probably the mammal most adapted for gliding, with a patagium that is as large as geometrically possible. They can glide as far a 70 m with minimal loss of height.
  • Sifaka and possibly some other primates (possible limited gliding/parachuting) . A number of primates have been suggested to have adaptations that allow limited gliding and/or parachuting; sifakas, indris, galagos and saki monkeys. Most notably the sifaka, a type of lemur, has thick hairs on its forearms that have been argued to provide drag, and a small membrane under its arms that has been suggested to provide lift by having aerofoil properties.[24] [7]
  • Cats and maybe others.[10] (very limited parachuting). If they fall cats spread their bodies to maximize drag, a very limited form of parachuting. Cats have an innate ‘righting reflex‘ that allows them to rotate their bodies so they fall feet first. Some other animals may show similar very limited parachuting. There are also anecdotal accounts of less limited parachuting, or even semi-gliding, in palm civets.[24]

[edit] Animals which parachute, glide, or fly (extinct)

Pterosaurs included the largest known flying animals

[edit] Reptiles

  • Extinct reptiles similar to Draco (gliding). There are a number of unrelated extinct lizard-like reptiles with similar “wings” to the Draco lizards. Icarosaurus, Daedalosaurus, Coelurosauravus, Weigeltosaurus, Mecistotrachelos,[25] and Kuehneosaurus. The largest of these, Kuehneosaurus, has a wingspan of 30 cm, and was estimated to be able to glide about 30 m.
  • Sharovipteryx (gliding). This strange reptile, sometimes proposed as a pterosaur ancestor, from the Upper Triassic of Kirghiia unusually had a membrane on its elongated hind limbs, as opposed to the forelimbs, which is much more usual. In some reconstructions they had webbing on the forelimbs and neck as well.[26]
  • Longisquama insignis (possibly gliding/parachuting). This small reptile may have had long paired feather-like scales on its back, however it has been more recently argued that the scales form just a single dorsal frill. If paired, they may have been used for parachuting.[27][28] “Everything you can make out is consistent with it being a small, tree-living, gliding animal, which is precisely the thing you’d expect birds to evolve out of,” says Larry Martin, senior curator at the Natural History Museum at the University of Kansas [8].
  • Pterosaurs (flying). Pterosaurs were the first flying vertebrates, and are generally agreed to have been sophisticated flyers. They had large wings formed by a patagium stretching from the torso to a dramatically lengthened fourth finger. There were hundreds of species, most of which are thought to have been intermittent flappers, and many soarers. The largest known flying animals are pterosaurs.

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