Animals that have evolved aerial locomotion
Greylag geese (
Anser anser
). Birds are one of only four taxonomic groups to have
evolved
powered
flight
.
A number of
animals
are capable of aerial locomotion, either by powered
flight
or by
gliding
. This trait has appeared by
evolution
many times, without any single common ancestor. Flight has evolved at least four times in separate animals:
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
and
reptiles
have also evolved this gliding flight ability, typically as a means of evading predators.
Types
[
edit
]
Animal aerial locomotion can be divided into two categories: powered and unpowered. In unpowered modes of locomotion, the animal uses aerodynamic 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 to climb or to maintain steady, level flight. Those who can find air that is rising faster than they are falling can gain altitude by
soaring
.
Unpowered
[
edit
]
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 descent. 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
[
edit
]
Powered
flight
has evolved at least four times: first in the
insects
, then in
pterosaurs
, next in
birds
, and last in
bats
. Studies on
theropod
dinosaurs do suggest multiple (at least 3) independent acquisitions of powered flight however,
[1]
[2]
and a recent study proposes independent acquisitions amidst the different bat clades as well.
[3]
Powered flight uses muscles to generate
aerodynamic force
, which allows the animal to produce lift and thrust. The animal may ascend without the aid of rising air.
Externally powered
[
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]
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
[
edit
]
Gliding and parachuting
[
edit
]
While gliding occurs independently from powered flight,
[4]
it has some ecological advantages of its own as it is the simplest form of flight.
[5]
Gliding is a very
energy
-efficient way of travelling from tree to tree. Although moving through the canopy running along the branches may be less energetically demanding, the faster transition between trees allows for greater foraging rates in a particular patch.
[6]
Glide ratios can be dependent on size and current behavior. Higher foraging rates are supported by low glide ratios as smaller foraging patches require less gliding time over shorter distances and greater amounts of food can be acquired in a shorter time period.
[6]
Low ratios are not as energy efficient as the higher ratios,
[5]
but 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.
[7]
Mammals tend to rely on lower glide ratios to increase the amount of time foraging for lower energy food.
[8]
An equilibrium glide, achieving a constant airspeed and glide angle, is harder to obtain as animal size increases. Larger animals need to glide from much higher heights and longer distances to make it energetically beneficial.
[9]
Gliding is also very suitable for predator avoidance, allowing for controlled targeted landings to safer areas.
[10]
[9]
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 India or 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.
[11]
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. Forest structure and distance between trees are influential in the development of gliding within varying species.
[8]
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 may be a factor.
[11]
In Australia, many mammals (and all mammalian gliders) possess, to some extent, prehensile tails. Globally, smaller gliding species tend to have feather-like tails and larger species have fur covered round bushy tails,
[10]
but smaller animals tend to rely on parachuting rather than developing gliding membranes.
[9]
The gliding membranes,
patagium
, are classified in the 4 groups of propatagium, digipatagium, plagiopatagium and uropatagium. These membranes consist of two tightly bounded layers of skin connected by muscles and connective tissue between the fore and hind limbs.
[10]
Powered flight evolution
[
edit
]
Analogous flying adaptions in vertebrates
:
Powered flight has evolved unambiguously only four times?
birds
,
bats
,
pterosaurs
, and
insects
(though see above for possible independent acquisitions within bird and bat groups). 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
.
[12]
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 evolved from tracheal gill structures and were used to catch the wind for small insects that live on the surface of the water, while another is that they evolved from paranotal lobes or leg structures and gradually progressed from parachuting, to gliding, to flight for originally arboreal insects.
[13]
Pterosaurs
were the next to evolve flight, approximately 228 million years ago. These reptiles were close relatives of the dinosaurs, 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,
Archaeopteryx
is 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 of 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). It may also have been a non-linear process, as several non-avian dinosaurs seem to have independently acquired powered flight.
[14]
[15]
Bats
are the most recent to evolve (about 60 million years ago), most likely from a fluttering ancestor,
[16]
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 area relative to their weight, which maximizes lift.
[17]
Soaring is very energetically efficient.
Biomechanics
[
edit
]
Gliding and parachuting
[
edit
]
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 tree nearby. Specifically in gliding mammals, there are 3 types of gliding paths respectively being S glide, J glide, and "straight-shaped" glides where species either gain altitude post launch then descend, rapidly decrease height before gliding, and maintaining a constant angled descent.
[10]
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. Successful flights for gliding animals are achieved through 5 steps: preparation, launch, glide, braking, and landing. Gliding species are better able to control themselves mid-air, with the tail acting as a rudder, making it capable to pull off banking movements or U-turns during flight.
[10]
During landing, arboreal mammals will extend their fore and hind limbs in front of itself to brace for landing and to trap air in order to maximize air resistance and lower impact speed.
[10]
Powered flight
[
edit
]
Large milkweed bug
flying
, repeated at one fifteenth speed.
Unlike most air vehicles, in which the objects that generate lift (wings) and thrust (engine or propeller) are separate and the wings remain 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.
[18]
There are two basic aerodynamic models of insect flight. Most insects use a method that creates a spiralling
leading edge
vortex
.
[19]
[20]
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.
[19]
[20]
Limits and extremes
[
edit
]
Flying and soaring
[
edit
]
- 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
pterosaur
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, 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,
[21]
but there is no firm evidence that any flying animal, not even the azhdarchid pterosaurs, got that large.
Comparison of
Quetzalcoatlus northropi
with a Cessna 172 light aircraft
- Smallest. There is no minimum size for getting airborne. Indeed, there are many bacteria floating in the atmosphere that constitute part of the
aeroplankton
.
[22]
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.
[
citation needed
]
The smallest flying invertebrate is a
fairyfly wasp
species,
Kikiki huna
, at 0.15 mm (0.0059 in) (150 μm).
[23]
- 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;
[24]
that measurement does not separate any contribution from wind speed, so the observations could be caused by strong
tailwinds
.
[25]
- 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).
[26]
- Highest flying. There are records of a
Ruppell's vulture
Gyps rueppelli
, a large vulture, being sucked into a jet engine 11,550 metres (37,890 ft) above
Cote d'Ivoire
in West Africa.
[27]
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).
[28]
Airborne flying squirrel.
Gliding and parachuting
[
edit
]
- 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 about 2.
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,
[29]
and thus fall just 1 meter 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.
[30]
[31]
[32]
Flying animals
[
edit
]
Extant
[
edit
]
A
bee
in flight.
- Pterygota
: The first of all animals to evolve flight, they are also the only invertebrates that have evolved flight. As they comprise almost all insects, the species are too numerous to list here.
Insect flight
is an active research field.
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
for birds that can soar as well as fly.
Townsends's big-eared bat
, (
Corynorhinus townsendii
) displaying the "hand wing"
- Bats
. There are approximately 1,240 bat species, representing about 20% of all classified mammal species.
[33]
Most bats are nocturnal and many feed on insects while flying at night, using
echolocation
to home in on their prey.
[34]
Extinct
[
edit
]
Pterosaurs
included the largest known flying animals
- 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.
Non-avian dinosaurs
[
edit
]
- Theropods
(gliding and 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
gui
,
Microraptor zhaoianus
, 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. A recent study indicates that flight may have been acquired independently in various different lineages
[2]
though it may have only evolved in theropods of the
Avialae
.
Gliding animals
[
edit
]
Extant
[
edit
]
Insects
[
edit
]
- Gliding bristletails
. 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
[35]
- Gliding ants
. 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.
[36]
Unique among gliding animals, Cephalotini and Pseudomyrmecinae ants glide abdomen first, the Forminicae however glide in the more conventional head first manner.
[37]
- 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
,
mantis
,
katydid
,
stick insect
and
true bug
.
Neon flying squid
- Flying squid
. Several oceanic
squids
of the family
Ommastrephidae
, such as the
Pacific flying squid
, will leap out of the water to escape predators, an adaptation similar to that of
flying fish
.
[38]
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.
[39]
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).
[40]
Band-winged flying fish
, with enlarged
pectoral fins
Fish
[
edit
]
- Flying fish
. 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).
[41]
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
[42]
). It has been suggested that the genus
Exocoetus
is on an evolutionary borderline between flight and gliding. It flaps its large
pectoral fins
while gliding, but does not use a power strike like flying animals.
[43]
It has been found that some flying fish can glide as effectively as some flying birds.
[44]
- live bearers
- Halfbeaks
. 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 separate hops.
[45]
- Trinidadian guppies
have been observed exhibiting a gliding response to escape predator's
[46]
[47]
[48]
- 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.
[49]
However, it is debated whether the freshwater butterfly fish can truly glide, Saidel et al. (2004) argue that it cannot.
- Freshwater hatchetfish
. In the wild, they have been observed jumping out of the water and gliding
[50]
(although reports of them achieving powered flight has been brought up many times
[51]
[52]
[53]
).
Illustration of
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
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.
- Rhacophoridae
flying frogs
. A number of the Rhacophoridae, such as
Wallace's flying frog
(
Rhacophorus nigropalmatus
), have adaptations for gliding, the main feature being enlarged toe membranes. For example, the
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
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
).
[54]
[55]
- Hylidae
flying frogs
. The other frog family that contains gliders.
[56]
Reptiles
[
edit
]
Several lizards and snakes are capable of gliding:
Gliding
Draco
lizard
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.
[41]
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 or parachuting.
[57]
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.
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.
[58]
- Lupersaurus
flying geckos
. A possible sister-taxon to
Ptychozoon
which has similar flaps and folds and also glides.
[58]
- Thecadactylus
flying geckos
. At least some species of
Thecadactylus
, such as
T. rapicauda
, are known to glide.
[58]
- 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.
Mammals
[
edit
]
Bats
are the only
freely flying mammals
.
[59]
A few other mammals can glide or parachute; the best known are
flying squirrels
and
flying lemurs
.
- Flying squirrels
(subfamily
Petauristinae
). There are more than 40 living species divided between 14 genera of flying
squirrel
. Flying squirrels are found in Asia (most species), North America (genus
Glaucomys
) and Europe (
Siberian flying squirrel
). They inhabit tropical, temperate, and
Subarctic
environments, with the
Glaucomys
preferring boreal and montane coniferous forests,
[60]
specifically landing on red spruce (
Picea rubens
) trees as landing sites; they are known to rapidly climb trees, but take some time to locate a good landing spot.
[61]
They tend to be
nocturnal
and are highly sensitive to light and noise.
[60]
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
). 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
). 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. They have the most developed propatagium out of any gliding mammal with a mean launch velocity of approximately 3.7 m/s;
[62]
the Mayan Colugo has been known to initiate glides without jumping.
[10]
- Sifaka
, a type of lemur, and possibly some other primates (possible limited gliding or parachuting). A number of primates have been suggested to have adaptations that allow limited gliding 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.
[63]
[64]
- Flying phalangers
or wrist-winged gliders (subfamily
Petaurinae
). Possums
[65]
[66]
[67]
[68]
[69]
[70]
[71]
[72]
[73]
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
and the
Biak glider
are the most common species. The lone species in the genus
Gymnobelideus
,
Leadbeater's possum
has only a vestigial gliding membrane.
- Greater glider
(
Petauroides volans
). The only species of the genus
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
Petaurinae
.
[74]
It has elongated limbs compared to its non-gliding relatives.
[10]
- Feather-tailed possums
(family
Acrobatidae
). 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.
Extinct
[
edit
]
Reptiles
[
edit
]
Life restoration of the
Weigeltisaurid
Weigeltisaurus jaekeli
from the Late Permian (259-252 million years ago). Weigeltisaurids represent the oldest known gliding vertebrates
Extinct reptiles similar to
Draco
. There are a number of unrelated extinct lizard-like reptiles with similar "wings" to the
Draco
lizards. These include the Late Permian
Weigeltisauridae
, the Triassic
Kuehneosauridae
and
Mecistotrachelos
,
[75]
and the Cretaceous lizard
Xianglong
. 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
. 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.
[76]
- Hypuronector
. This bizarre drepanosaur displays limb proportions, particularly the elongated forelimbs, that are consistent with a flying or gliding animal with
patagia
.
[77]
Non-avian dinosaurs
[
edit
]
Life restoration of
Yi qi
a gliding
scansoriopterygid
dinosaur from the Middle Jurassic of China.
Scansoriopterygidae
is unique among 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.
Fish
[
edit
]
Volaticotherids
predate bats as mammalian aeronauts by at least 110 million years
Mammals
[
edit
]
See also
[
edit
]
References
[
edit
]
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Further reading
[
edit
]
- Davenport, J. (1994). "How and why do flying fish fly?".
Reviews in Fish Biology and Fisheries
.
40
(2): 184?214.
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:
1994RFBF....4..184D
.
doi
:
10.1007/BF00044128
.
S2CID
34720887
.
- 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
(1): 63?72.
Bibcode
:
2004EnvBF..71...63S
.
doi
:
10.1023/b:ebfi.0000043153.38418.cd
.
S2CID
11856131
.
- Xu, Xing; Zhou, Zhonghe; Wang, Xiaolin; Kuang, Xuewen; Zhang, Fucheng; Du, Xiangke (2003).
"Four-winged dinosaurs from China"
(PDF)
.
Nature
.
421
(6921): 335?340.
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:
2003Natur.421..335X
.
doi
:
10.1038/nature01342
.
PMID
12540892
.
S2CID
1160118
.
- Schiøtz, A.; Vosloe, H. (1959). "The gliding flight of Holaspis guentheri Gray, a west-African lacertid".
Copeia
.
1959
(3): 259?260.
doi
:
10.2307/1440407
.
JSTOR
1440407
.
- 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
(2): 155?163.
doi
:
10.1017/s0968047002000171
.
S2CID
49552361
.
- McGuire, J. A. (2003). "Allometric Prediction of Locomotor Performance: An Example from Southeast Asian Flying Lizards".
The American Naturalist
.
161
(2): 337?349.
doi
:
10.1086/346085
.
PMID
12675377
.
S2CID
29494470
.
- Demes, B.; Forchap, E.; Herwig, H. (1991). "They seem to glide. Are there aerodynamic effects in leaping prosimian primates?".
Zeitschrift fur Morphologie und Anthropologie
.
78
(3): 373?385.
doi
:
10.1127/zma/78/1991/373
.
PMID
1909482
.
- The Pterosaurs: From Deep Time by David Unwin
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