Flying and gliding animals
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 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
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
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 than true flyers.
Powered flight
Externally powered
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.
Evolution and ecology
Gliding and parachuting
While gliding occurs independently from powered flight, it has some ecological advantages of its own as it is the simplest form of flight. 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. 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. Low ratios are not as energy efficient as the higher ratios, 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. Mammals tend to rely on lower glide ratios to increase the amount of time foraging for lower energy food. 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. Gliding is also very suitable for predator avoidance, allowing for controlled targeted landings to safer areas. In contrast to flight, gliding has evolved independently many times ; 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. Additionally, a variety of gliding vertebrates are found in Africa, a family of hylids 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 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. 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 in Asian forests may be a factor. In Australia, many mammals 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, but smaller animals tend to rely on parachuting rather than developing gliding membranes. 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.
Powered flight evolution
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 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, 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.
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. However, they spanned a large range of sizes, down to a 250 mm 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 or a "ground up" origin. It may also have been a non-linear process, as several non-avian dinosaurs seem to have independently acquired powered flight.
Bats are the most recent to evolve, 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 area relative to their weight, which maximizes lift.