A number of animals have evolved aerial locomotion, either by powered
flight or by gliding.
Flying and gliding animals
Flying and gliding animals (volant animals) have
evolved separately many times, without any single ancestor.
evolved at least four times, in the insects, pterosaurs, birds, and
bats. Gliding has evolved on many more occasions. Usually the
development is to aid canopy animals in getting from tree to tree,
although there are other possibilities. Gliding, in particular, has
evolved among rainforest animals, especially in the rainforests in
Asia (most especially Borneo) where the trees are tall and widely
spaced. Several species of aquatic animals, and a few amphibians have
also evolved to acquire this gliding flight ability, typically as a
means of evading predators.
1.2 Powered flight
1.3 Externally powered
Evolution and ecology
2.1 Gliding and parachuting
2.2 Powered flight
3.1 Gliding and parachuting
3.2 Powered flight
4 Limits and extremes
5 Extant flying and gliding animals
6 Extinct flying and gliding animals
6.2 Non-avian dinosaurs
7 See also
9 Further reading
10 External links
Animal aerial locomotion can be divided into two categories—powered
and unpowered. In unpowered modes of locomotion, the animal uses
aerodynamics forces exerted on the body due to wind or falling through
the air. In powered flight, the animal uses muscular power to generate
aerodynamic forces. Animals using unpowered aerial locomotion cannot
maintain altitude and speed due to unopposed drag, while animals using
powered flight can maintain steady, level flight as long as their
muscles are capable of doing so.
These modes of locomotion typically require an animal start from a
raised location, converting that potential energy into kinetic energy
and using aerodynamic forces to control trajectory and angle of
Energy is continually lost to drag without being replaced,
thus these methods of locomotion have limited range and duration.
Falling: decreasing altitude under the force of gravity, using no
adaptations to increase drag or provide lift.
Parachuting: falling at an angle greater than 45° from the horizontal
with adaptations to increase drag forces. Very small animals may be
carried up by the wind. Some gliding animals may use their gliding
membranes for drag rather than lift, to safely descend.
Gliding flight: falling at an angle less than 45° from the horizontal
with lift from adapted aerofoil membranes. This allows slowly falling
directed horizontal movement, with streamlining to decrease drag
forces for aerofoil efficiency and often with some maneuverability in
air. Gliding animals have a lower aspect ratio (wing length/breadth)
than true flyers.
Powered flight has evolved only four times (first in insects, then in
pterosaurs, birds and bats). It uses muscular power to generate
aerodynamic forces and to replace energy lost to drag.
Flapping: moving wings to produce lift and thrust. May ascend without
the aid of the wind, as opposed to gliders and parachuters.
Ballooning and soaring are not powered by muscle, but rather by
external aerodynamic sources of energy: the wind and rising thermals,
respectively. Both can continue as long as the source of external
power is present. Soaring is typically only seen in species capable of
powered flight, as it requires extremely large wings.
Ballooning: being carried up into the air from the aerodynamic effect
on long strands of silk in the wind. Certain silk-producing
arthropods, mostly small or young spiders, secrete a special
light-weight gossamer silk for ballooning, sometimes traveling great
distances at high altitude.
Soaring: gliding in rising or otherwise moving air that requires
specific physiological and morphological adaptations that can sustain
the animal aloft without flapping its wings. The rising air is due to
thermals, ridge lift or other meteorological features. Under the right
conditions, soaring creates a gain of altitude without expending
energy. Large wingspans are needed for efficient soaring.
Many species will use multiple of these modes at various times; a hawk
will use powered flight to rise, then soar on thermals, then descend
via free-fall to catch its prey.
Evolution and ecology
Gliding and parachuting
While gliding occurs independently from powered flight, it has some
ecological advantages of its own. Gliding is a very energy-efficient
way of travelling from tree to tree. An argument made is that many
gliding animals eat low energy foods such as leaves and are restricted
to gliding because of this, whereas flying animals eat more high
energy foods such as fruits, nectar, and insects. In contrast to
flight, gliding has evolved independently many times (more than a
dozen times among extant vertebrates), however these groups have not
radiated nearly as much as have groups of flying animals.
Worldwide, the distribution of gliding animals is uneven as most
inhabit rain forests in Southeast Asia. (Despite seemingly suitable
rain forest habitats, few gliders are found in
New Guinea and
none in Madagascar.) Additionally, a variety of gliding vertebrates
are found in Africa, a family of hylids (flying frogs) lives in South
America and several species of gliding squirrels are found in the
forests of northern
Asia and North America. Various factors produce
these disparities. In the forests of Southeast Asia, the dominant
canopy trees (usually dipterocarps) are taller than the canopy trees
of the other forests. A higher start provides a competitive advantage
of further glides and farther travel. Gliding predators may more
efficiently search for prey. The lower abundance of insect and small
vertebrate prey for carnivorous animals (such as lizards) in Asian
forests ( In Australia, many mammals (and all mammalian gliders)
possess, to some extent, prehensile tails.
Analogous flying adaptions in vertebrates:
Powered flight has evolved unambiguously only four times—birds,
bats, pterosaurs, and insects. In contrast to gliding, which has
evolved more frequently but typically gives rise to only a handful of
species, all three extant groups of powered flyers have a huge number
of species, suggesting that flight is a very successful strategy once
evolved. Bats, after rodents, have the most species of any mammalian
order, about 20% of all mammalian species.
Birds have the most
species of any class of terrestrial vertebrates. Finally, insects
(most of which fly at some point in their life cycle) have more
species than all other animal groups combined.
The evolution of flight is one of the most striking and demanding in
animal evolution, and has attracted the attention of many prominent
scientists and generated many theories. Additionally, because flying
animals tend to be small and have a low mass (both of which increase
the surface-area-to-mass ratio), they tend to fossilize infrequently
and poorly compared to the larger, heavier-boned terrestrial species
they share habitat with. Fossils of flying animals tend to be confined
to exceptional fossil deposits formed under highly specific
circumstances, resulting in a generally poor fossil record, and a
particular lack of transitional forms. Furthermore, as fossils do not
preserve behavior or muscle, it can be difficult to discriminate
between a poor flyer and a good glider.
Insects were the first to evolve flight, approximately 350 million
years ago. The developmental origin of the insect wing remains in
dispute, as does the purpose prior to true flight. One suggestion is
that wings initially were used to catch the wind for small insects
that live on the surface of the water, while another is that they
functioned in parachuting, then gliding, then flight for originally
Pterosaurs were the next to evolve flight, approximately 228 million
years ago. These reptiles were close relatives of the dinosaurs (and
sometimes mistakenly considered dinosaurs by laymen), and reached
enormous sizes, with some of the last forms being the largest flying
animals ever to inhabit the Earth, having wingspans of over 9.1 m
(30 ft). However, they spanned a large range of sizes, down to a
250 mm (10 in) wingspan in Nemicolopterus.
Birds have an extensive fossil record, along with many forms
documenting both their evolution from small theropod dinosaurs and the
numerous bird-like forms of theropod which did not survive the mass
extinction at the end of the Cretaceous. Indeed,
arguably the most famous transitional fossil in the world, both due to
its mix of reptilian and avian anatomy and the luck of being
discovered only two years after Darwin's publication of On the Origin
of Species. However, the ecology and this transition is considerably
more contentious, with various scientists supporting either a "trees
down" origin (in which an arboreal ancestor evolved gliding, then
flight) or a "ground up" origin (in which a fast-running terrestrial
ancestor used wings for a speed boost and to help catch prey).
Bats are the most recent to evolve (about 60 million years ago), most
likely from a fluttering ancestor, though their poor fossil record
has hindered more detailed study.
Only a few animals are known to have specialised in soaring: the
larger of the extinct pterosaurs, and some large birds. Powered flight
is very energetically expensive for large animals, but for soaring
their size is an advantage, as it allows them a low wing loading, that
is a large wing areas relative to their weight, which maximizes
lift. Soaring is very energetically efficient.
Gliding and parachuting
During a free-fall with no aerodynamic forces, the object accelerates
due to gravity, resulting in increasing velocity as the object
descends. During parachuting, animals use the aerodynamic forces on
their body to counteract the force or gravity. Any object moving
through air experiences a drag force that is proportion to surface
area and to velocity squared, and this force will partially counter
the force of gravity, slowing the animal's descent to a safer speed.
If this drag is oriented at an angle to the vertical, the animal's
trajectory will gradually become more horizontal, and it will cover
horizontal as well as vertical distance. Smaller adjustments can allow
turning or other maneuvers. This can allow a parachuting animal to
move from a high location on one tree to a lower location on another
During gliding, lift plays an increased role. Like drag, lift is
proportional to velocity squared. Gliding animals will typically leap
or drop from high locations such as trees, just as in parachuting, and
as gravitational acceleration increases their speed, the aerodynamic
forces also increase. Because the animal can utilize lift and drag to
generate greater aerodynamic force, it can glide at a shallower angle
than parachuting animals, allowing it to cover greater horizontal
distance in the same loss of altitude, and reach trees further away.
Unlike most air vehicles, in which the objects that generate lift
(wings) and thrust (engine/propeller) are separate and the wings
remained fixed, flying animals use their wings to generate both lift
and thrust by moving them relative to the body. This has made the
flight of organisms considerably harder to understand than that of
vehicles, as it involves varying speeds, angles, orientations, areas,
and flow patterns over the wings.
A bird or bat flying through the air at a constant speed moves its
wings up and down (usually with some fore-aft movement as well).
Because the animal is in motion, there is some airflow relative to its
body which, combined with the velocity of its wings, generates a
faster airflow moving over the wing. This will generate lift force
vector pointing forwards and upwards, and a drag force vector pointing
rearwards and upwards. The upwards components of these counteract
gravity, keeping the body in the air, while the forward component
provides thrust to counteract both the drag from the wing and from the
body as a whole.
Pterosaur flight likely worked in a similar manner,
though no living pterosaurs remain for study.
Insect flight is considerably different, due to their small size,
rigid wings, and other anatomical differences. Turbulence and vortices
play a much larger role in insect flight, making it even more complex
and difficult to study than the flight of vertebrates. There are
two basic aerodynamic models of insect flight. Most insects use a
method that creates a spiralling leading edge vortex. Some very
small insects use the fling-and-clap or Weis-Fogh mechanism in which
the wings clap together above the insect's body and then fling apart.
As they fling open, the air gets sucked in and creates a vortex over
each wing. This bound vortex then moves across the wing and, in the
clap, acts as the starting vortex for the other wing. Circulation and
lift are increased, at the price of wear and tear on the wings.
Limits and extremes
This section needs additional citations for verification. Please help
improve this article by adding citations to reliable sources.
Unsourced material may be challenged and removed. (October 2012)
(Learn how and when to remove this template message)
Largest. The largest known flying animal was formerly thought to be
Pteranodon, a pterosaur with a wingspan of up to 7.5 metres
(25 ft). However, the more recently discovered azhdarchid
Quetzalcoatlus is much larger, with estimates of the
wingspan ranging from 9 to 12 metres (30 to 39 ft). Some other
recently discovered azhdarchid pterosaur species, such as
Hatzegopteryx, may have also wingspans of a similar size or even
slightly larger. Although it is widely thought that Quetzalcoatlus
reached the size limit of a flying animal, it should be noted that the
same was once said of Pteranodon. The heaviest living flying animals
are the kori bustard and the great bustard with males reaching 21
kilograms (46 lb). The wandering albatross has the greatest
wingspan of any living flying animal at 3.63 metres (11.9 ft).
Among living animals which fly over land, the
Andean condor and the
marabou stork have the largest wingspan at 3.2 metres (10 ft).
Studies have shown that it is physically possible for
flying animals to reach 18-metre (59 ft) wingspans, but there is
no firm evidence that any flying animal, not even the azhdarchid
pterosaurs, got that large.
Smallest. There is no real minimum size for getting airborne. Indeed,
there are many bacteria floating in the atmosphere that constitute
part of the aeroplankton. However, to move about under one's own power
and not be overly affected by the wind requires a certain amount of
size. The smallest flying vertebrates are the bee hummingbird and the
bumblebee bat, both of which may weigh less than 2 grams
(0.071 oz). They are thought to represent the lower size limit
for endotherm flight.
Fastest. The fastest of all known flying animals is the peregrine
falcon, which when diving travels at 300 kilometres per hour
(190 mph) or faster. The fastest animal in flapping horizontal
flight may be the Mexican free-tailed bat, said to attain about 160
kilometres per hour (99 mph) based on ground speed by an aircraft
tracking device; that measurement does not separate any
contribution from wind speed, so the observations could be caused by
Slowest. Most flying animals need to travel forward to stay aloft.
However, some creatures can stay in the same spot, known as hovering,
either by rapidly flapping the wings, as do hummingbirds, hoverflies,
dragonflies, and some others, or carefully using thermals, as do some
birds of prey. The slowest flying non-hovering bird recorded is the
American woodcock, at 8 kilometres per hour (5.0 mph).
Highest flying. There are records of a
Rüppell's vulture Gyps
rueppelli, a large vulture, being sucked into a jet engine 11,550
metres (37,890 ft) above
Côte d'Ivoire in West Africa. The
animal that flies highest most regularly is the bar-headed goose Anser
indicus, which migrates directly over the
Himalayas between its
nesting grounds in
Tibet and its winter quarters in India. They are
sometimes seen flying well above the peak of
Mount Everest at 8,848
metres (29,029 ft).
Most efficient glider. This can be taken as the animal that moves most
horizontal distance per metre fallen. Flying squirrels are known to
glide up to 200 metres (660 ft), but have measured glide ratio of
Flying fish have been observed to glide for hundreds of
metres on the drafts on the edge of waves with only their initial leap
from the water to provide height, but may be obtaining additional lift
from wave motion. On the other hand, albatrosses have measured
lift/drag ratios of 20, and thus fall just 1 meter (foot) for
every 20 in still air.
Most maneuverable glider. Many gliding animals have some ability to
turn, but which is the most maneuverable is difficult to assess. Even
paradise tree snakes, Chinese gliding frogs, and gliding ants have
been observed as having considerable capacity to turn in the air.
Extant flying and gliding animals
A bee in flight.
Insects (flying). 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 flight is an active research
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 
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
Camponotus). All arboreal dolichoderines and non-cephalotine
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. Unique among gliding
Pseudomyrmecinae ants glide abdomen first,
the Forminicae however glide in the more conventional head first
Gliding immature insects. The wingless immature stages of some insect
species that have wings as adults may also show a capacity to glide.
These include some species of cockroach, mantid, katydid, stick insect
and true bug. 
Spiders. Although typically flightless some may engage in aerial
locomotion as described below.
Ballooning 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.
Gliding spiders (gliding). Some species of arboreal spider of the
Selenops can glide back to the trunk of a tree should they fall.
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. Smaller squids will fly
in shoals, and have been observed to cover distances as long as 50
metres (160 ft). 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
providing thrust even after leaving the water. This may make flying
squid the only animals with jet-propelled aerial locomotion. The
neon flying squid has been observed to glide for distances over 30
metres (100 ft), at speeds of up to 11.2 metres per second
(37 ft/s) .
Band-winged flying fish, with 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
centimetres (18 in) but most species measure less than 30 cm
(12 in) 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 kilometres per hour (37 mph). The
glides are usually up to 30–50 metres (100–160 ft) 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 ). It has been suggested that the genus
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. It has been found that some
flying fish can glide as effectively as some flying birds.
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 metres (160 ft) in two
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. 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[citation
Wallace's flying frog
Wallace's flying frog in Alfred Russel Wallace's 1869
book The Malay Archipelago
Gliding has evolved independently in two families of tree frogs, the
Rhacophoridae and the
New World Hylidae. Within each lineage
there are a range of gliding abilities from non-gliding, to
parachuting, to full gliding.
Rhacophoridae flying frogs (gliding). A number of the Rhacophoridae,
Wallace's flying frog
Wallace's flying frog (Rhacophorus nigropalmatus), have
adaptations for gliding, the main feature being enlarged toe
membranes. For example, the
Malayan flying frog
Malayan flying frog Rhacophorus prominanus
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 flying frog
Chinese flying frog Rhacophorus 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
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.
Several lizards and snakes are capable of gliding:
Draco lizards. 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 metres (200 ft) and over this distance they lose only 10
metres (30 ft) in height. 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. 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.
Ptychozoon flying geckos. 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.
Lupersaurus flying geckos. A possible sister-taxon to
has similar flaps and folds and also glides.
Thecadactylus flying geckos. At least some species of Thecadactylus,
such as T. rapicauda, are known to glide.
Cosymbotus flying gecko. Similar adaptations to
Ptychozoon are found
in the two species of the gecko genus Cosymbotus.
Chrysopelea snakes. 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 metres
(330 ft) and make 90 degree turns.
Birds are a successful group of flying vertebrate.
Birds (flying, soaring) — Most of the approximately 10,000 living
species can fly (flightless birds are the exception).
Bird flight is
one of the most studied forms of aerial locomotion in animals. See
List of soaring birds
List of soaring birds for birds that can soar as well as fly.
Bats are the only mammal with flapping or powered flight. A few other
mammals glide or parachute; the best known are flying squirrels and
Bats (flying). There are approximately 1,240 bat species, representing
about 20% of all classified mammal species.
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 metres (660 ft).
Anomalures or scaly-tailed flying squirrels (family Anomaluridae)
(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 have gliding membranes between their front and hind legs.
The genus Idiurus contains two particularly small species known as
flying mice, but similarly they are not true mice.
Colugos or "flying lemurs" (order Dermoptera) (gliding). There are two
species of colugo. Despite their common name, colugos are not lemurs;
true lemurs are primates. Molecular evidence suggests that colugos are
a sister group to primates; however, some mammalogists 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 as 70 metres
(230 ft) with minimal loss of height.
Sifaka, a type of lemur, 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.
Flying phalangers or wrist-winged gliders (subfamily Petaurinae)
(gliding). Possums found in
Australia, and New Guinea. The gliding membranes are hardly noticeable
until they jump. On jumping, the animal extends all four legs and
stretches the loose folds of skin. The subfamily contains seven
species. Of the six species in the genus Petaurus, the sugar glider
Biak glider are the most common species. The lone species in
the genus Gymnobelideus,
Leadbeater's possum has only a vestigial
Greater glider (
Petauroides volans) (gliding). The only species of the
Petauroides of the family Pseudocheiridae. This marsupial is
found in Australia, and was originally classed with the flying
phalangers, but is now recognised as separate. Its flying membrane
only extends to the elbow, rather than to the wrist as in
Feather-tailed possums (family Acrobatidae) (gliding). This family of
marsupials contains two genera, each with one species. The feathertail
glider (Acrobates pygmaeus), found in
Australia is the size of a very
small mouse and is the smallest mammalian glider. The feathertail
possum (Distoechurus pennatus) is found in New Guinea, but does not
glide. Both species have a stiff-haired feather-like tail.
Townsends's big-eared bat, (Corynorhinus townsendii) displaying the
Extinct flying and gliding animals
Pterosaurs included the largest known flying animals
Extinct reptiles similar to Draco (gliding). There are a number of
unrelated extinct lizard-like reptiles with similar "wings" to the
Draco lizards. Icarosaurus, Coelurosauravus, Weigeltisaurus,
Mecistotrachelos, and Kuehneosaurus. The largest of these,
Kuehneosaurus, has a wingspan of 30 centimetres (12 in), and was
estimated to be able to glide about 30 metres (100 ft).
Sharovipterygidae (gliding). These strange reptiles from the Upper
Triassic of Kyrgyzstan and
Poland unusually had a membrane on their
elongated hind limbs, extending their otherwise normal,
flying-squirrel-like patagia significantly. The forelimbs are in
contrast much smaller.
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. "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.
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.
Hypuronector (gliding). This bizarre drepanosaur displays limb
proportions, particularly the elongated forelimbs, that are consistent
with a flying or gliding animal with patagia.
Theropods (gliding/flying). There were several species of theropod
dinosaur thought to be capable of gliding or flying, that are not
classified as birds (though they are closely related). Some species
Microraptor zhaoianus, Cryptovolans pauli, and
Changyuraptor) have been found that were fully feathered on all four
limbs, giving them four 'wings' that they are believed to have used
for gliding or flying. One species,
Deinonychus antirrhopus, may
display partial volancy, with the young being capable of flight while
the adults are flightless, a characteristic also seen in some modern
birds like the
Horned coot and the Flying steamer duck.
Yi is unique among gliding dinosaurs for the development of membranous
wings, unlike the feathered airfoils of other theropods. Much like
modern anomalures it developed a bony rod to help support the wing,
albeit on the wrist and not the elbow.
Thoracopteridae (gliding) is a lineage of
Triassic flying fish-like
Perleidiformes, having converted their pectoral and pelvic fins into
broad wings very similar to those of their modern counterparts. The
Potanichthys is the oldest member of this clade, as
well as the earliest aerial vertebrate known, suggesting that these
fish began exploring aerial niches soon after the Permian-Triassic
Volaticotherids predate bats as mammalian aeronauts by at least 110
Volaticotherium antiquum (gliding). A gliding eutriconodont, long
considered the earliest gliding mammal until the discovery of
contemporary gliding haramiyidans. It lived around 164 million years
ago and used a fur-covered skin membrane to glide through the air.
The closely related
Argentoconodon is also thought to have been able
to glide, based on postcranial similarities; it lived around 165
million years ago.
Several species of extinct bat have been found, like Icaronycteris,
Palaeochiropteryx, and Onychonycteris.
A gliding metatherian (possibly a marsupial) is known from the
Paleocene of Itaboraí, Brazil.
The haramiyidans Vilevolodon, Xianshou,
Arboroharamiya had extensive patagia, highly convergent with those of
Flying mythological creatures
Organisms at high altitude
^ "Life in the Rainforest". Archived from the original on 2006-07-09.
Retrieved 15 April 2006.
^ a b Corlett, Richard T.; Primack, Richard B. (2011). Tropical rain
forests : an ecological and biogeographical comparison (2nd ed.).
Chichester: Wiley-Blackwell. pp. 197, 200.
^ Simmons, N.B.; D.E. Wilson, D.C. Reeder (2005).
the World: A Taxonomic and Geographic Reference. Baltimore, MD: Johns
Hopkins University Press. pp. 312–529.
^ Kaplan, Matt (2011). "Ancient bats got in a flap over food". Nature.
Vertebrate Flight". Retrieved 15 April 2006.
^ Wang, Shizhao; Zhang, Xing; He, Guowei; Liu, Tianshu (Sep 2013).
"Lift Enhancement by Dynamically Changing
Wingspan in Forward Flapping
Flight". Physics of Fluids. 26: 061903. arXiv:1309.2726 .
^ a b Wang, Z. Jane (2005). "DISSECTING INSECT FLIGHT" (pdf). Annual
Review of Fluid Mechanics. Annual Reviews. 37: 183.
^ a b Sane, Sanjay P. (2003). "The aerodynamics of insect flight"
(PDF). The Journal of Experimental Biology. 206 (23): 4191–4208.
doi:10.1242/jeb.00663. PMID 14581590.
^ McCracken, Gary F.; Safi, Kamran; Kunz, Thomas H.; Dechmann, Dina K.
N.; Swartz, Sharon M.; Wikelski, Martin (9 November 2016). "Airplane
tracking documents the fastest flight speeds recorded for bats". Royal
Society Open Science. 3: 160398. Bibcode:2016RSOS....360398M.
^ Photopoulos, Julianna (9 November 2016). "Speedy bat flies at
160km/h, smashing bird speed record". New Scientist. Retrieved 11
November 2016. But not everyone is convinced. Graham Taylor at the
University of Oxford says that errors in estimating bat speed by
measuring the distance moved between successive positions could be
huge. “So I think it would be premature to knock birds off their
pedestal as nature's fastest fliers just yet,” he says."These bats
are indeed flying very fast at times, but this is based on their
ground speed," says Anders Hedenström at the University of Lund in
Sweden. "Since they did not measure winds at the place and time where
the bats are flying, one can therefore not exclude that the top speeds
are not bats flying in a gust."
^ Yanoviak, SP; Kaspari, M; Dudley, R (2009). "Gliding hexapods and
the origins of insect aerial behaviour". Biology Letters. 5 (4):
510–2. doi:10.1098/rsbl.2009.0029. PMC 2781901 .
^ Yanoviak, S. P.; Dudley, R.; Kaspari, M. (2005). "Directed aerial
descent in canopy ants". Nature. 433 (7026): 624–626.
^ "Scientist Discovers
Rainforest Ants That Glide". Newswise.
Retrieved 15 April 2006.
^ Packard, A. (1972). "Cephalopods and fish: the limits of
convergence". Biological Reviews. 47 (2): 241–307.
^ Maciá, Silvia; Robinson, Michael P.; Craze, Paul; Dalton, Robert;
Thomas, James D. (2004). "New observations on airborne jet propulsion
(flight) in squid, with a review of previous reports". Journal of
Molluscan Studies. 70 (3): 297–299.
^ a b Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of
Curious and Unusual Animals, Greenwood Press.
^ BBC NEWS Science/Nature Fast flying fish glides by ferry
Vertebrate Flight: gliding and parachuting". Retrieved 15 April
Flying fish perform as well as some birds - Los Angeles Times
^ Marshall, N.B. (1965) The Life of Fishes. London: Weidenfield and
Nicolson. 402 pp.
^ Berra, Tim M. (2001). Freshwater
Fish Distribution. San Diego:
Academic Press. ISBN 0-12-093156-7
^ McKay, M. G. (2001). "Aerodynamic stability and maneuverability of
the gliding frog Polypedates dennysi". Journal of Experimental
Biology. 204 (16): 2817–2826.
^ Emerson, Sharon B.; Koehl, M. A. R. (1990). "The interaction of
behavioral and morphological change in the evolution of a novel
locomotor type: "flying" frogs". Evolution. 44 (8): 1931–1946.
doi:10.2307/2409604. JSTOR 2409604.
^ Mendelson, Joseph R; Savage, Jay M; Griffith, Edgardo; Ross, Heidi;
Kubicki, Brian; Gagliardo, Ronald (2008). "Spectacular new gliding
species of Ecnomiohyla (Anura: Hylidae) from Central Panama". Journal
of Herpetology. 42 (4): 750–759. doi:10.1670/08-025R1.1.
^ Tiny lizard falls like a feather
^ a b c Ptychozoon: the geckos that glide with flaps and fringes
(gekkotans part VIII) – Tetrapod Zoology
^ Tudge, Colin (2000). The Variety of Life. Oxford University Press.
^ Darren Naish: Tetrapod Zoology: Literally, flying lemurs (and not
^ Literally, flying lemurs (and not dermopterans) – Tetrapod Zoology
Archived 16 August 2010 at the Wayback Machine.
^ Gliding Possums — Environment, New South Wales Government
^ Cronin, Leonard — "Key Guide to Australian Mammals", published by
Reed Books Pty. Ltd., Sydney, 1991 ISBN 0-7301-0355-2
^ van der Beld, John — "Nature of
Australia — A portrait of the
island continent", co-published by William Collins Pty. Ltd. and ABC
Enterprises for the Australian Broadcasting Corporation, Sydney, 1988
(revised edition 1992), ISBN 0-7333-0241-6
^ Russell, Rupert — "Spotlight on Possums", published by University
of Queensland Press, St. Lucia, Queensland, 1980,
^ Troughton, Ellis — "Furred Animals of Australia", published by
Angus and Robertson (Publishers) Pty. Ltd, Sydney, in 1941 (revised
edition 1973), ISBN 0-207-12256-3
^ Morcombe, Michael & Irene — "
Mammals of Australia", published
by Australian Universities Press Pty. Ltd, Sydney, 1974,
^ Ride, W. D. L. — "A Guide to the Native
Mammals of Australia",
published by Oxford University Press, Melbourne, 1970, ISBN 0 19
^ Serventy, Vincent — "Wildlife of Australia", published by Thomas
Nelson (Australia) Ltd., Melbourne, 1968 (revised edition 1977),
^ Serventy, Vincent (editor) — "Australia's Wildlife Heritage",
published by Paul Hamlyn Pty. Ltd., Sydney, 1975
^ Myers, Phil. "Family Pseudocheiridae". Retrieved 15 April
^ Ancient Gliding Reptile Discovered LiveScience
^ Dzik, J.; Sulej, Tomasz (2016). "An early Late
reptile with a bony pectoral shield and gracile appendages" (PDF).
Acta Palaeontologica Polonica. 64 (4): 805–823.
^ Stauth, David (2000). "Ancient feathered animal challenges
dinosaur-bird link". Archived from the original on 11 June 2004.
Retrieved 15 April 2006.
^ "Controversial Fossil Claimed to Sink Dinosaur-
Bird Link". Archived
from the original on 2006-06-30. Retrieved 15 April 2006.
Dinosaur Profs Worlds Apart on Link to Birds
^ Renesto, S., Spielmann, J. A., Lucas, S. G., & Spagnoli, G. T.
(2010). The taxonomy and paleobiology of the Late Triassic
(Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida:
Archosauromorpha: Drepanosauromorpha): Bulletin 46 (Vol. 46). New
Mexico Museum of Natural History and Science.
^ BBC NEWS Science/Nature Earliest flying mammal discovered
^ Gaetano, L.C.; Rougier, G.W. (2011). "New materials of
Argentoconodon fariasorum (Mammaliaformes, Triconodontidae) from the
Jurassic of Argentina and its bearing on triconodont phylogeny".
Vertebrate Paleontology. 31 (4): 829–843.
^ Simmons, N.B.; Seymour, K.L.; Habersetzer, J.; Gunnell, G.F.
(February 14, 2008). "Primitive Early Eocene bat from Wyoming and the
evolution of flight and echolocation". Nature. 451 (7180): 818–822.
^ Szalay, FS, Sargis, EJ, and Stafford, BJ (2000) Small marsupial
glider from the
Paleocene of Itaboraí, Brazil. Journal of Vertebrate
Paleontology 20 Supplement: 73A. Presented at the Meeting of the
^ Meng, Qing-Jin; Grossnickle, David M.; Di, Liu; Zhang, Yu-Guang;
Neander, April I.; Ji, Qiang; Luo, Zhe-Xi (2017). "New gliding
mammaliaforms from the Jurassic". Nature. 548: 291–296.
Davenport, J. (1994). "How and why do flying fish fly?". Reviews in
Fish Biology and Fisheries. 40: 184–214.
Saidel, W.M.; Strain, G.F.; Fornari, S.K. (2004). "Characterization of
the aerial escape response of the African butterfly fish, Pantodon
buchholzi Peters". Environmental Biology of Fishes. 71: 63–72.
Xu, Xing; Zhou, Zhonghe; Wang, Xiaolin; Kuang, Xuewen; Zhang, Fucheng;
Du, Xiangke (2003). "Four-winged dinosaurs from China". Nature. 421
(6921): 335–340. Bibcode:2003Natur.421..335X.
doi:10.1038/nature01342. PMID 12540892.
Schiøtz, A.; Vosloe, H. (1959). "The gliding flight of Holaspis
guentheri Gray, a west-African lacertid". Copeia. 1959: 259–260.
Arnold, E. N. (2002). "Holaspis, a lizard that glided by accident:
mosaics of cooption and adaptation in a tropical forest lacertid
(Reptilia, Lacertidae. )". Bulletin of the Natural History Museum.
Zoology Series. 68: 155–163. doi:10.1017/s0968047002000171.
McGuire, J. A. (2003). "Allometric Prediction of Locomotor
Performance: An Example from Southeast Asian Flying Lizards". The
American Naturalist. 161: 337–349. doi:10.1086/346085.
Demes, B.; Forchap, E.; Herwig, H. (1991). "They seem to glide. Are
there aerodynamic effects in leaping prosimian primates?". Zeitschrift
für Morphologie und Anthropologie. 78: 373–385.
The Pterosaurs: From Deep Time by David Unwin
Wikimedia Commons has media related to
Canopy Locomotion from
Mongabay online magazine
Learn the Secrets of
Flight Exhibit at UCMP
Insect flight, photographs of flying insects — Rolf Nagels
Map of Life - "Gliding mammals" — University of Cambridge
Fins, limbs and wings
Fin and flipper locomotion
Flying and gliding animals
Evolution of fish
Evolution of tetrapods
Evolution of birds
Origin of birds
Origin of avian flight
Evolution of cetaceans
Tradeoffs for locomotion in air and water
Animal sexual behaviour
Animal welfare science
Bee learning and communication
Emotion in animals
Pain in animals
Tool use by animals
Karl von Frisch
William Homan Thorpe
Jakob von Uexküll
E. O. Wilson
Association for the Study of
International Society for Applied Ethology