Spiracles () are openings on the surface of some

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

s, which usually lead to

respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...

The spiracle is a small hole behind each eye that opens to the mouth in some

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...

. In the

jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present ( cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclosto ...

, the first

gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

opening immediately behind the mouth is essentially similar to the other gill opening. With the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

of the

jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...

in the early jawed vertebrates, this gill slit was caught between the forward gill-rod (now functioning as the jaw) and the next rod, the hyomandibular bone, supporting the jaw hinge and anchoring the jaw to the

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...

proper. The gill opening was closed off from below, the remaining opening was small and hole-like, and is termed a spiracle. In many species of sharks and all rays the spiracle is responsible for the intake of water into the

buccal space The buccal space (also termed the buccinator space) is a fascial space of the head and neck (sometimes also termed fascial tissue spaces or tissue spaces). It is a potential space in the cheek, and is paired on each side. The buccal space is supe ...

before being expelled from the gills. The spiracle is often located towards the top of the animal allowing breathing even while the animal is mostly buried under sediments. As sharks adapted a faster moving lifestyle some became obligate ram ventilators, breathing exclusively by forcing water through their gills by swimming, among these are

requiem shark Requiem sharks are sharks of the family Carcharhinidae in the order Carcharhiniformes. They are migratory, live-bearing sharks of warm seas (sometimes of brackish or fresh water) and include such species as the tiger shark, bull shark, le ...

s and

hammerhead shark The hammerhead sharks are a group of sharks that form the family Sphyrnidae, so named for the unusual and distinctive structure of their heads, which are flattened and laterally extended into a "hammer" shape called a cephalofoil. Most hammerhe ...

s which have lost their spiracles In

elasmobranchs Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...

(sharks and rays) the spiracle bears a small

pseudobranch The pseudobranch, also pseudobranchia is the reduced first gill arch of a fish (on the inner surface of the opercle, near the junction of the preopercle) as well as a reduced "false" gill in some gastropods. In teleost fish, the pseudobranchs are m ...

that resembles a gill in structure, but only receives blood already oxygenated by the true gills. The function of the pseudobranch is unknown, but it is believed that it supplies highly oxygenated blood to the optic choroid and retina and may have baroreceptor (pressure) and thermoregulation functions. It may also be a site of oxygen chemoreception. Chimaeras lack spiracles, using gill opercula for

buccal pumping Buccal pumping is "breathing with one's cheeks": a method of ventilation used in respiration in which the animal moves the floor of its mouth in a rhythmic manner that is externally apparent.Brainerd, E. L. (1999). New perspectives on the evolutio ...

instead.

Bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...

have similar gill opercula but the basalmost ray-finned fish

bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pu ...

s use their spiracles for inhaling air into their lungs, this leads to speculation this may be the original air breathing mechanism ancestral to all bony fish and

tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...

Coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...

s have closed off spiracles which may be a product of their deepwater lifestyle and loss of air breathing lungs. Bichirs as a whole may more closely resemble the common ancestor of lobe-finned fish and bony fish as a whole than coelacanths due to their deepwater adaptations.

Acipenseriformes Acipenseriformes is an order of basal ray-finned fishes that includes living and fossil sturgeons and paddlefishes (Acipenseroidei), as well as the extinct families Chondrosteidae and Peipiaosteidae. They are the second earliest div ...

including

sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...

s and

paddlefish Paddlefish (family Polyodontidae) are a family of ray-finned fish belonging to order Acipenseriformes, and one of two living groups of the order alongside sturgeons (Acipenseridae). They are distinguished from other fish by their titular elongla ...

have small seemingly vestigial spiracles much like coelacanths further reduced in

Holostei Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by a single living species, the bowfin ('' Amia calva''), as well as the Ginglymodi, the sole living representatives being the gars ...

and completely absent in

Teleostei Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest class (biology), infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of a ...

, the clade containing 96% of all extant species of fish. In tetrapods the spiracle seems to have developed first into the

otic notch Otic notches are invaginations in the posterior margin of the skull roof, one behind each orbit. Otic notches are one of the features lost in the evolution of amniotes from their tetrapod ancestors. The notches have been interpreted as part of a ...

of early tetrapods where it was still used in respiration and incapable of sensing sound, and then into the ear of modern tetrapods which by the Eustachian tube remains connected to the buccal cavity. The spiracle is still found in all cartilaginous fish except

s, and chimaeras, and is found in some primitive bony fishes (

coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...

and

s). It is also seen as an

in the skull of the extinct

labyrinthodonts "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consi ...

, and is thought to be associated with the

ear An ear is the organ that enables hearing and, in mammals, body balance using the vestibular system. In mammals, the ear is usually described as having three parts—the outer ear, the middle ear and the inner ear. The outer ear consists of ...

opening in

amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...

s and

frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...

s. Romer, A.S. (1949): ''The Vertebrate Body.'' W.B. Saunders, Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970) Blowholes in cetaceans are also sometimes referred to as spiracles, but they are not homologous with the spiracles of fish, having instead developed from the trachea. In cetaceans and other mammals, the organs homologous with the spiracles of fish are the ears.

References

{{reflist Respiratory system Vertebrate anatomy