Amia Calva
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The ruddy bowfin (''Amia calva'') is a
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
, being one of only two surviving species of the
Halecomorphi Halecomorphi is a taxon of Actinopterygii, ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin (''Amia calva'') and eyespot bowfin (''Amia ocellicauda''), but the group contains many extinct species i ...
, a group of fish that first appeared during the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
, around 250 million years ago. The bowfin is often considered a "
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
" because they have retained some morphological characteristics of their early ancestors. It is one of two species in the genus ''Amia,'' along with ''
Amia ocellicauda ''Amia ocellicauda'', the eyespot bowfin or emerald bowfin, is a species of Amia (fish), bowfin native to North America. Originally described by John Richardson (naturalist), John Richardson from Lake Huron in 1836, it was synonymized with ''Amia ...
'', the eyespot bowfin. The closest living relatives of bowfins are
gar Gars are an ancient group of ray-finned fish in the family Lepisosteidae. They comprise seven living species of fish in two genera that inhabit fresh, brackish, and occasionally marine waters of eastern North America, Central America and Cuba ...
s, with the two groups being united in the clade
Holostei Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, '' Amia'' with two species, the bowfins (''Amia calva'' and '' Amia ocellicauda''), as well as the Gin ...
. Bowfins are
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer o ...
freshwater
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
s, commonly found throughout much of the eastern United States, and in southern
Ontario Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
and
Quebec Quebec is Canada's List of Canadian provinces and territories by area, largest province by area. Located in Central Canada, the province shares borders with the provinces of Ontario to the west, Newfoundland and Labrador to the northeast, ...
. Fossil deposits indicate
Amiiformes The Amiiformes order (biology), order of fish has only two extant taxa, extant species, the bowfins: ''Amia calva'' and ''Amia ocellicauda'', the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater syste ...
were once widespread in both freshwater and marine environments across North and South America, Europe, Asia, and Africa. Now, their range is limited to much of the eastern United States and adjacent southern Canada, including the
drainage basin A drainage basin is an area of land in which all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
s of the
Mississippi River The Mississippi River is the main stem, primary river of the largest drainage basin in the United States. It is the second-longest river in the United States, behind only the Missouri River, Missouri. From its traditional source of Lake Ita ...
,
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
, and various rivers exiting in the Eastern Seaboard or
Gulf of Mexico The Gulf of Mexico () is an oceanic basin and a marginal sea of the Atlantic Ocean, mostly surrounded by the North American continent. It is bounded on the northeast, north, and northwest by the Gulf Coast of the United States; on the southw ...
. Their preferred habitat includes vegetated sloughs, lowland rivers and lakes, swamps, and backwater areas; they are also occasionally found in brackish water. They are stalking, ambush predators known to move into the shallows at night to prey on fish and aquatic invertebrates such as crawfish, mollusks, and aquatic insects. Like gars, bowfin are bimodal breathers—they have the capacity to breathe both water and air. Their
gills 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 ...
exchange gases in the water allowing them to breathe, but they also have a
gas bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift via swimming, ...
that serves to maintain buoyancy, and also allows them to breathe air by means of a small
pneumatic duct The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift via swimming, wh ...
connected from the foregut to the gas bladder. They can break the surface to gulp air, which allows them to survive conditions of aquatic hypoxia that would be lethal to most other species. The bowfin is long-lived, with age up to 33 years reported.


Morphology

The typical length of a bowfin is ; females typically grow to , males to . They can reach in length, and weigh . Young of the year typically grow to by October. Females tend to grow larger than males. The body of the bowfin is elongated and cylindrical, with the sides and back
olive The olive, botanical name ''Olea europaea'' ("European olive"), is a species of Subtropics, subtropical evergreen tree in the Family (biology), family Oleaceae. Originating in Anatolia, Asia Minor, it is abundant throughout the Mediterranean ...
to brown in color, often with vertical bars and dark reticulations or another camouflaged pattern. The dorsal fin has horizontal bars, and the caudal fin has irregular vertical bars. The underside is white or cream, and the paired fins and anal fin are bright green. During larval stage, hatchlings from about total length are black and tadpole-like in appearance. At approximately total length they have been described as looking like miniature
placoderms Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos plate' and δέρμα 'dermaskin') are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Dev ...
. They grow quickly, and typically leave the nest within 4 to 6 weeks after hatching. Young males have a black eyespot on the base of the tail (
caudal peduncle Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only ...
) that is commonly encircled by an orange-yellowish border, while the female's is black, if present at all. It is thought the purpose of the eyespot is to confuse predators, deflecting attacks away from the head of the fish to its tail, which affords the bowfin an opportunity to escape predation. The bowfin is so named for its long, undulating
dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...
consisting of 145 to 250 rays that runs from the middle of the back to the base of the tail. The skull of the bowfin is made of two layers of skull, the
dermatocranium The dermatocranium is the portion of the cranium that is composed of dermal bone, as opposed to the endocranium and splanchnocranium, which are composed of endochondral bone. The dermatocranium comprises the skull roof, the facial skeleton (usu ...
and the
chondrocranium The chondrocranium (or ''cartilaginous neurocranium'') is the primitive cartilaginous skeletal structure of the fetal skull that grows to envelop the rapidly growing embryonic brain.Salentijn, L. ''Biology of Mineralized Tissues: Prenatal Skul ...
. The chondrocranium layer cannot be seen because it is located below the dermal bones. The bowfin skull is made up of 28 fused bones, which compose the dermatocranium. The roof of the mouth is made up of three bones, the ectopterygoid, the palantine, and the vomer. They have two sets of teeth, including one set of larger sharp teeth coming out of the
mandibular In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
ry bones to grasp and control the prey. The other set of teeth, located posteriorly and connected to the
hyomandibula The hyomandibula, commonly referred to as hyomandibular one(, from , "upsilon-shaped" (υ), and Latin: mandibula, "jawbone"), is a set of bones that is found in the hyoid region in most fishes. It usually plays a role in suspending the jaws ...
r bone, is made up of pharyngeal tooth patches, which are used for sorting out nutrients and grinding down larger pieces of food. '' ''Another three bones make up the lower jaw: the dentary, the angular, and the surangular. The cranial surface of the skull is made up of the nasals, the antorbital, the lacrimal, the parietal, the intertemporal, the post parietal, the supratemporal, the extra scapular, the post temporal, and the opercular. The entirety of the skull is attached to the girdle through another set of bones. Bowfin are often referred to as "
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
s" or " primitive fish" because they retained some of the primitive characters common to their ancestors, including a modified (rounded externally)
heterocercal Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only ...
caudal fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only ...
, a highly vascularized
gas bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift via swimming, ...
lung, vestiges of a
spiral valve A spiral valve or scroll valve is the corkscrew-shaped lower portion of the intestine of some sharks, Acipenseriformes (sturgeon and paddlefish), rays, skates, bichirs, Lepisosteiformes (gars), and lungfishes. A modification of the ileum, th ...
, and a bony gular plate. The bony gular plate is located underneath the head on the exterior of the lower jaw between the two sides of the lower jaw bone. Other distinguishing characteristics include long, sharp teeth, and two protruding tube-like nostrils. Unlike all of the most primitive
actinopterygians Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin ...
, the scales of bowfin differ in that they are not ganoid scales, rather they are large, single-layered
cycloid In geometry, a cycloid is the curve traced by a point on a circle as it Rolling, rolls along a Line (geometry), straight line without slipping. A cycloid is a specific form of trochoid and is an example of a roulette (curve), roulette, a curve g ...
scales closer in similarity to more derived
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
s.


Fish similar in appearance

Northern snakehead The northern snakehead (''Channa argus'') is a species of snakehead fish native to temperate East Asia, in China, Russia, North Korea, and South Korea. Their natural range goes from the Amur River watershed in Siberia and Manchuria down to Hainan ...
s (''Channa argus'') are commonly mistaken for bowfin because of similarities in appearance, most noticeably their elongated, cylindrical shape, and long dorsal fin that runs along their backs. Northern snakeheads are
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
fish native to the rivers and estuaries of China, Russia, and Korea that have been introduced and become established in parts of North America. However, unlike bowfin which are native to North America, the northern snakehead is considered an
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
and environmentally harmful there. Some contrasting differences in bowfin include a black eyespot on their caudal peduncle, a tan and olive coloration, a shorter anal fin, a more rounded head, pelvic fins at a greater distance from the pectoral fins than in the northern snakehead and the presence of the gular plate on the ventral side of the lower jaw. Another noticeable difference is that bowfin scales do not continue uniformly from their body to their head. Bowfin heads are smooth and free of scales, whereas the northern snakehead has scales that uniformly continue from their body through to their head. The
burbot The burbot (''Lota lota''), also known as bubbot, mariah, loche, cusk, freshwater cod, freshwater ling, freshwater cusk, the lawyer, coney-fish, lingcod, or eelpout, is a species of coldwater ray-finned fish native to the subarctic regions of ...
(''Lota lota''), a
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
y fish native to streams and lakes of North America and Eurasia, is also commonly mistaken for bowfin. Burbots can be distinguished by their flat head and chin barbel, long anal fin, and pelvic fins situated beneath the pectoral fins.


Bowfin body-shape evolution and development

The first fish lacked jaws and used negative pressure to suck their food in through their mouths. The jaw in the bowfin is a result of their evolutionary need to be able to catch and eat bigger and more nutritious prey. As a result of being able to gather more nutrients, Bowfin are able to live a more active lifestyle. The jaw of a bowfin has several adaptations. The maxilla and premaxilla are fused and the posterior chondrocranium articulates with the vertebra which allows the jaw freedom to rotate. The suspensorium includes several bones and articulates with the snout, brain case, and the mandible. When the jaw opens
epaxial muscles In adult vertebrates, trunk muscles can be broadly divided into hypaxial muscles, which lie ventral to the horizontal septum of the vertebrae and epaxial muscles, which lie dorsal to the septum. Hypaxial muscles include some vertebral muscles, the ...
lift the chondrocranium, which is attached to the upper jaw, while adductor muscles act to close the lower jaw. This ability to open and close the jaw helps the bowfin to be an active predator that can catch bigger prey and digest them. The
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
in bowfin is ossified and in comparison to earlier fish, the
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the individual shops are all owned by individual franchisees. The chain has three ...
are the major support for the body, whereas in earlier fish the notochord was the main form of support. In bowfin neural spines and ribs also increase in prominence, an evolutionary aspect that helps provide additional support and stabilize unpaired fins. The evolution of the vertebral column allows the bowfin to withstand lateral bending that puts the column under compression without breaking. This, in turn, allows the bowfin to have more controlled and powerful movements, in comparison to fish that had only a notochord. The bowfin has a rounded heterocercal tail that resembles a homocercal tail. This type of tail gives the body a streamlined shape which allows the bowfin to improve its swimming ability by reducing drag. These types of tails are common in fish with gas bladders, because the bladder supplies the fish with natural buoyancy. The bowfin is a member of
actinopterygii Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class (biology), class of Osteichthyes, bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built ...
which means that the pectoral girdle is partly
endochondral Endochondral ossification is one of the two essential pathways by which bone tissue is produced during fetal development and bone repair of the mammalian skeletal system, the other pathway being intramembranous ossification. Both endochondral an ...
but mostly
dermal bone A dermal bone or investing bone or membrane bone is a bony structure derived from intramembranous ossification forming components of the vertebrate skeleton, including much of the skull, jaws, gill covers, shoulder girdle, fin rays ( lepidotrich ...
. In this group of fish the fins function to maneuver, brake, and for slight positional adjustments. The pectoral girdle of the bowfin has six parts. The post temporal, supracleithrum, postcleithrum, cleithrum, scapulacoracoid, and the clavicle make up the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
. The pectoral girdle is attached to the skull. The paired pectoral and pelvic fins of fish are homologous with the limbs of tetrapods.


Physiology

Bowfin are
physostome Physostomes are fishes that have a pneumatic duct connecting the gas bladder to the alimentary canal. This allows the gas bladder to be filled or emptied via the mouth. This not only allows the fish to fill their bladder by gulping air, but al ...
s, meaning they have a small "
pneumatic duct The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift via swimming, wh ...
" that connects their swim bladders to their digestive tract. This allows them, like
lungfish Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
, to "breathe" in two ways: they can extract oxygen from the water when breathing through their
gills 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 ...
, but can also break the water's surface to breathe or gulp air through the
pneumatic duct The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift via swimming, wh ...
. When performing low-level physical activity, bowfin obtain more than half of their oxygen from breathing air. The fish have two distinct air-breathing mechanisms used to ventilate the gas bladder. Air breathing ''type I'' is consistent with the action of exhale / inhale exchange, stimulated by either air or water hypoxia, to regulate gas exchange; ''type II'' air breaths are inhalation alone, which is believed to regulate gas bladder volume, to control
buoyancy Buoyancy (), or upthrust, is the force exerted by a fluid opposing the weight of a partially or fully immersed object (which may be also be a parcel of fluid). In a column of fluid, pressure increases with depth as a result of the weight of t ...
. Bimodal respiration helps bowfin survive and maintain their metabolic rate in
hypoxic Hypoxia means a lower than normal level of oxygen, and may refer to: Reduced or insufficient oxygen * Hypoxia (environmental), abnormally low oxygen content of a specific environment ** Hypoxia in fish, responses of fish to hypoxia * Hypoxia (medi ...
(low-oxygen) conditions. Bowfin air breathe more frequently when they are in darkness, and correspondingly more active. Bowfin blood can adapt to warm,
acid An acid is a molecule or ion capable of either donating a proton (i.e. Hydron, hydrogen cation, H+), known as a Brønsted–Lowry acid–base theory, Brønsted–Lowry acid, or forming a covalent bond with an electron pair, known as a Lewis ...
ic waters. The fish becomes inactive in waters below ; at this temperature they breathe almost no air; however, with increasing temperature their air breathing increases. Their preferred temperature range is between , with the temperature of maximum activity. Air breathing is at a maximum in the range . Bowfin do not use central chemoreceptor regulation for respiration control. Experiments manipulating the oxygen content, carbon dioxide content, and pH of bowfin extradural fluid did not affect breathing rate, heart rate, or blood pressure pointing to a lack of central chemoreceptor regulation. Instead, bowfin respiratory patterns respond to water oxygen content and water temperature, as water temperatures play a role in oxygen content. In the lab, bowfin showed an increase in the breathing rate when the temperatures were raised above 10°C. Bowfin also showed an increase in breathing rate when exposed to lower oxygen levels in the water. Herpetologist W. T. Neill reported in 1950 that he unearthed a bowfin aestivating (in a dormant state) in a chamber below the ground surface, in diameter, from a river. It was further noted that flood levels had previously reached the area, and receded. It is not unusual for riverine species like bowfin to move into backwaters with flood currents, and become trapped when water levels recede. While aestivation is anecdotally documented by multiple researchers, laboratory experiments have suggested instead that bowfin are physiologically incapable of surviving more than three to five days of air exposure. However, no field manipulation has been performed. Regardless of the lack of evidence confirming the bowfin's ability to aestivate, it has been noted that bowfin can survive prolonged conditions of exposure to air because they have the ability to breathe air. Their gill filaments and
lamellae Lamella (: lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * Lame ...
are rigid in structure which helps prevent the lamellae from collapsing and aids gas exchange even during air exposure.


Evolution and phylogeny

Competing
hypotheses A hypothesis (: hypotheses) is a proposed explanation for a phenomenon. A scientific method, scientific hypothesis must be based on observations and make a testable and reproducible prediction about reality, in a process beginning with an educ ...
and debates continue over the evolution of ''Amia'' and relatives, including their relationship among basal extant teleosts, and organization of
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s. Bowfin are the last remaining member of
Halecomorphi Halecomorphi is a taxon of Actinopterygii, ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin (''Amia calva'') and eyespot bowfin (''Amia ocellicauda''), but the group contains many extinct species i ...
, a group that includes many extinct species in several families. Halecomorphs were generally accepted as the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to
Teleostei Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
but not without question. While a halecostome pattern of
neopterygian Neopterygii (from Greek νέος ''neos'' 'new' and πτέρυξ ''pteryx'' 'fin') is a subclass of ray-finned fish (Actinopterygii). Neopterygii includes the Holostei and the Teleostei, of which the latter comprise the vast majority of extant f ...
clades was produced in morphology-based analyses of extant
actinopterygian Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin ...
s, a different result was produced with fossil taxa which showed a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
Holostei. Monophyletic Holostei were also recovered by at least two
nuclear gene A nuclear gene is a gene whose DNA sequence is located within the cell nucleus of a eukaryotic organism. These genes are distinguished from extranuclear genes, such as those found in the genomes of mitochondria and chloroplasts, which reside ou ...
analyses, in an independent study of fossil and extant fish, and in an analysis of ultraconserved genomic elements. The extant ray-finned fish of the subclass Actinopterygii include 42 orders, 431 families and over 23,000 species. They are currently classified into two infraclasses,
Chondrostei Chondrostei is a subclass of non- neopterygian ray-finned fish. While the term originally referred to the paraphyletic grouping of all non-neopterygian ray-finned fish, it was redefined by Patterson in 1982 to be a clade comprising the Acipenser ...
(
holosteans Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, '' Amia'' with two species, the bowfins (''Amia calva'' and ''Amia ocellicauda''), as well as the Ginglymo ...
) and
Neopterygii Neopterygii (from Greek νέος ''neos'' 'new' and πτέρυξ ''pteryx'' 'fin') is a subclass of ray-finned fish (Actinopterygii). Neopterygii includes the Holostei and the Teleostei, of which the latter comprise the vast majority of extant ...
(teleost fishes). Sturgeons, paddlefish, bichirs and reed fish compose the thirty-eight species of chondrosteans, and are considered relict species. Included in the over 23,000 species of neopterygians are eight relict species comprising gars and the bowfin.


Infraclass Neopterygii

Neopterygians are the second major occurrence in the evolution of ray-finned fish and today include the majority of modern bony fish. They are distinguished from their earlier ancestors by major changes to the jaws, shape of the skull, and tail. They are divided into three divisions: * Division 1. Order
Lepisosteiformes Lepisosteiformes is an order of ray-finned fish and the only living members of the clade Ginglymodi. Its only extant representatives are the gar (family Lepisosteidae), and it is defined as all members of Ginglymodi that are more closely rela ...
 – the relict gars which include extant species of
gar Gars are an ancient group of ray-finned fish in the family Lepisosteidae. They comprise seven living species of fish in two genera that inhabit fresh, brackish, and occasionally marine waters of eastern North America, Central America and Cuba ...
s that first appeared in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. * Division 2. Order
Amiiformes The Amiiformes order (biology), order of fish has only two extant taxa, extant species, the bowfins: ''Amia calva'' and ''Amia ocellicauda'', the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater syste ...
 – the relict bowfin, (
halecomorphi Halecomorphi is a taxon of Actinopterygii, ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin (''Amia calva'') and eyespot bowfin (''Amia ocellicauda''), but the group contains many extinct species i ...
ds), the only extant species in the order
Amiiformes The Amiiformes order (biology), order of fish has only two extant taxa, extant species, the bowfins: ''Amia calva'' and ''Amia ocellicauda'', the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater syste ...
which date back to the
Triassic period The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is the ...
. * Division 3. Division Teleostei – the
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of
Teleostei Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
from which modern fish arose, including most of the
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
we are familiar with today.


Species

The following is a species list


Genome evolution

The bowfin
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
contains an intact
ParaHox The ParaHox gene cluster is an array of homeobox genes (involved in morphogenesis, the regulation of patterns of anatomical development) from the Gsx, Xlox ( Pdx) and Cdx gene families. Regulatory gene cluster These genes were first shown to ...
gene cluster A gene cluster is a group of two or more genes found within an organism's DNA that encode similar peptide, polypeptides or proteins which collectively share a generalized function and are often located within a few thousand base pairs of each othe ...
, similar to the
bichir Bichirs and the reedfish comprise Polypteridae , a family (biology), family of archaic Actinopterygii, ray-finned fishes and the only family in the order (biology), order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The D ...
and to most other
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. This is in contrast, however, with
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish, which have a fragmented ParaHox cluster, probably because of a
whole genome duplication Paleopolyploidy is the result of genome duplications which occurred at least several million years ago (MYA). Such an event could either double the genome of a single species ( autopolyploidy) or combine those of two species ( allopolyploidy). ...
event in their lineage. The presence of an intact ParaHox gene cluster suggests that bowfin ancestors separated from other fish before the
last common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...
of all
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
s appeared. Bowfin are thus possibly a better model to study vertebrate genome organization than common teleost
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s such as
zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...
.


Feeding behavior

Bowfin are stalking, ambush predators that customarily move into the shallows at night to prey on fish, amphibians, and aquatic invertebrates such as crawfish, other crustaceans, mollusks, and aquatic insects. Young bowfin feed mostly on small
crustaceans Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of Arthropod, arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquat ...
, while adults are mostly
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
, but also known to be opportunistic. Some common examples of prey include frogs, bass, other bowfin, dragonflies, sunfish, crawfish, etc. Bowfin are remarkably agile, can move quickly through the water, and they have a voracious appetite. Their undulating dorsal fin propels them silently through the water while stalking their prey. The attack is straightforward and swift with a movement that lasts approximately 0.075 seconds. There were also some studies regarding the capacity of the bowfin to survive without food. In 1916, a female bowfin was starved for twenty months. It was the longest period that any vertebrate had been without food, as far the writer was aware during the observation. Some independent studies focus on the bowfin's ability to use organic material as a source of food and studied the structure of the
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
. They concluded that it did not benefit from the organic material in the water because the gill rakers were short with blunt processes and a short space between them. Even bacteria could enter and exit through the gill easily. Its structure alone indicated that the ''Amia'' do not use microorganisms as a source of food.


Distribution and habitat

Fossil deposits indicate amiiforms included freshwater and marine species that were once widely distributed in North America, South America, Eurasia and Africa. Today, bowfin (''Amia calva'') are the only remaining species in the order Amiiformes; they are
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer o ...
freshwater
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
s, and their range is restricted to freshwater environments in North America, including much of the eastern
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
and adjacent southern Canada from the
St. Lawrence River The St. Lawrence River (, ) is a large international river in the middle latitudes of North America connecting the Great Lakes to the North Atlantic Ocean. Its waters flow in a northeasterly direction from Lake Ontario to the Gulf of St. Lawren ...
and
Lake Champlain Lake Champlain ( ; , ) is a natural freshwater lake in North America. It mostly lies between the U.S. states of New York (state), New York and Vermont, but also extends north into the Canadian province of Quebec. The cities of Burlington, Ve ...
drainage of southern
Ontario Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
and
Quebec Quebec is Canada's List of Canadian provinces and territories by area, largest province by area. Located in Central Canada, the province shares borders with the provinces of Ontario to the west, Newfoundland and Labrador to the northeast, ...
westward around the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
in southern Ontario into Minnesota. Historically, their distribution in North America included the
drainage basin A drainage basin is an area of land in which all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
s of the
Mississippi River The Mississippi River is the main stem, primary river of the largest drainage basin in the United States. It is the second-longest river in the United States, behind only the Missouri River, Missouri. From its traditional source of Lake Ita ...
from Quebec to northern Minnesota, the St. Lawrence-Great Lakes, including
Georgian Bay The Georgian Bay () is a large bay of Lake Huron, in the Laurentia bioregion. It is located entirely within the borders of Ontario, Canada. The main body of the bay lies east of the Bruce Peninsula and Manitoulin Island. To its northwest is t ...
,
Lake Nipissing Lake Nipissing (; , ) is a lake in the Provinces and territories of Canada, Canadian province of Ontario. It has a surface area of , a mean elevation of above sea level, and is located between the Ottawa River and Georgian Bay. Lake Nipissing i ...
and Simcoe, Ontario, south to the
Gulf of Mexico The Gulf of Mexico () is an oceanic basin and a marginal sea of the Atlantic Ocean, mostly surrounded by the North American continent. It is bounded on the northeast, north, and northwest by the Gulf Coast of the United States; on the southw ...
; Atlantic and
Gulf Coastal Plain The Gulf Coastal Plain extends around the Gulf of Mexico in the Southern United States and eastern Mexico. This coastal plain reaches from the Florida Panhandle, southwest Georgia (U.S. state), Georgia, the southern two-thirds of Alabama, over m ...
from the
Susquehanna River The Susquehanna River ( ; Unami language, Lenape: ) is a major river located in the Mid-Atlantic (United States), Mid-Atlantic region of the United States, crossing three lower Northeastern United States, Northeast states (New York, Pennsylvani ...
drainage in southeastern
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
to the
Colorado River The Colorado River () is one of the principal rivers (along with the Rio Grande) in the Southwestern United States and in northern Mexico. The river, the List of longest rivers of the United States (by main stem), 5th longest in the United St ...
in
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
.


Stocking

Research from the late 1800s to the 1980s suggests a trend of intentional stockings of non-indigenous fish into ponds, lakes and rivers in the United States. At that time, little was known about environmental impacts, or long-term effects of new species establishment and spread as a result of "fish rescue and transfer" efforts, or the importance of nongame fish to the ecological balance of aquatic ecosystems. Introductions of bowfin to areas they were considered a non-indigenous species included various lakes, rivers and drainages in Connecticut, Delaware, Georgia, Illinois, Iowa, Kansas, Kentucky, Maryland, Massachusetts, Minnesota, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Virginia, West Virginia, and Wisconsin. Many of the introductions were intentional stockings; however, there is no way to positively determine distribution resulting from flood transfers, or other inadvertent migrations. Bowfin are typically piscivorous, but as an introduced species are capable of being voracious predators that pose a threat to native fish and their prey.


Preferred habitat

Bowfin prefer vegetated sloughs, lowland rivers and lakes, swamps, backwater areas, and are occasionally found in brackish water. They are well camouflaged, and not easy to spot in slow water with abundant vegetation. They often seek shelter under roots, and submerged logs. Oxygen-poor environments can be tolerated because of their ability to breathe air.


Life cycle

Bowfin spawn in the spring or early summer, typically between April and June, more commonly at night in abundantly vegetated, clear shallow water in weed beds over sand bars, and also under stumps, logs, and bushes. Optimum temperatures for nesting and spawning range between . The males construct circular nests in fibrous root mats, clearing away leaves and stems. Depending on the density of surrounding vegetation there may be a tunnel-like entrance at one side. The diameter of the nests commonly range between , at a water depth of . During spawning season, the fins and underside of male bowfin often change in color to a bright lime green. The courtship/spawning sequence lasts one to three hours, and can repeat up to five times. Courtship begins when a female approaches the nest. The ritual consists of intermittent nose bites, nudges, and chasing behavior by the male until the female becomes receptive, at which time the pair lie side by side in the nest. She deposits her eggs while he shakes his fins in a vibratory movement, and releases his milt for fertilization to occur. A male often has eggs from more than one female in his nest, and a single female often spawns in several nests. Females vacate the nest after spawning, leaving the male behind to protect the eggs during the eight to ten days of incubation. A nest may contain 2,000 to 5,000 eggs, possibly more.
Fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the capability to produc ...
is usually related to size of the fish, so it is not unusual for the roe of a large gravid female to contain over 55,000 eggs. Bowfin eggs are adhesive, and will attach to aquatic vegetation, roots, gravel, and sand. After hatching, larval bowfin do not swim actively in search of food. During the seven to nine days required for yolk-sac absorption, they attach to vegetation by means of an adhesive organ on their snout, and remain protected by the parent male bowfin. Bowfin aggressively protect their spawn from the first day of incubation to a month or so after the eggs have hatched. When the fry are able to swim and forage on their own, they will form a school and leave the nest accompanied by the parent male bowfin who slowly circles them to prevent separation. Bowfin reach sexually maturity at two to three years of age. They can live up to 33 years in the wild, and 30 years in captivity. Bowfin may live decades at adult size.


Diseases

A common
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
of bowfin is the
anchor worm ''Lernaea'' is a genus of copepod crustaceans whose members are commonly called ''anchor worms'' and are parasitic on freshwater fishes. Life cycle Anchor worms mate during the last free-swimming (copepodid) stage of development. After mating, ...
(''Lernaea''). These small crustaceans infest the skin and bases of fins, with consequences ranging from slowed growth to death. The mollusk '' Megalonaias gigantea'' lays eggs in the bowfin gills, that are then externally fertilized by sperm passing in the water flow. The small
glochidia The glochidium (plural glochidia) is a microscopic larval stage of some freshwater mussels, aquatic bivalve mollusks in the families Unionidae and Margaritiferidae, the river mussels and European freshwater pearl mussels. These larvae are t ...
larvae then hatch and develop in the gill tubes. Bowfin with
liver cancer Liver cancer, also known as hepatic cancer, primary hepatic cancer, or primary hepatic malignancy, is cancer that starts in the liver. Liver cancer can be primary in which the cancer starts in the liver, or it can be liver metastasis, or secondar ...
and with fatal
leukemia Leukemia ( also spelled leukaemia; pronounced ) is a group of blood cancers that usually begin in the bone marrow and produce high numbers of abnormal blood cells. These blood cells are not fully developed and are called ''blasts'' or '' ...
have been reported.


Utilization

As a sport fish, bowfin are not considered desirable to many anglers. They were once considered a nuisance fish by anglers and early biologists who believed the bowfin's predatory nature was harmful to sport fish populations. As a result, efforts were taken to reduce their numbers. Research has since proven otherwise, and that knowledge together with a better understanding of maintaining overall balance of ecosystems, regulations were introduced to help protect and maintain viable populations of bowfin. Bowfin are strong fighters, a prized trait in game fish. However, they do have a jaw full of sharp teeth which requires careful handling. The current tackle record is Bowfin were once considered to have little commercial value because of its poor-tasting meat which has been referred to as "soft, bland-tasting and of poor texture". However, it is considered quite palatable if cleaned properly and
smoked Smoking is the process of flavoring, browning, cooking, or preserving food, particularly meat, fish and tea, by exposing it to smoke from burning or smoldering material, most often wood. In Europe, alder is the traditional smoking wood, but ...
, or prepared fried, blackened, used in courtbouillion, or in fishballs or fishcakes. Over the years, global efforts have imposed strict regulations on the international trade of caviar, particularly on the harvest of sturgeons from the Caspian Sea where the highly prized
caviar Caviar or caviare is a food consisting of salt-cured roe of the family Acipenseridae. Caviar is considered a delicacy and is eaten as a garnish or spread. Traditionally, the term caviar refers only to roe from wild sturgeon in the Caspi ...
from the
beluga sturgeon The beluga (), also known as the beluga sturgeon or great sturgeon (''Huso huso''), is a species of anadromous fish in the sturgeon family (Acipenseridae) of the order Acipenseriformes. It is found primarily in the Caspian and Black Sea basins, ...
originates. The bans imposed on Caspian sturgeons have created lucrative markets for affordable substitutes in the United States including
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 elongated rost ...
, bowfin, and various species of
sturgeon Sturgeon (from Old English ultimately from Proto-Indo-European language, Proto-Indo-European *''str̥(Hx)yón''-) is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the ...
. In
Louisiana Louisiana ( ; ; ) is a state in the Deep South and South Central regions of the United States. It borders Texas to the west, Arkansas to the north, and Mississippi to the east. Of the 50 U.S. states, it ranks 31st in area and 25 ...
, bowfin are harvested in the wild, and cultured commercially in hatcheries for their meat and roe. The roe is processed into caviar, and sold as "Cajun caviar", or marketed under the trade name "Choupiquet Royale".


Accumulation of toxic substances

In some areas of the United States where aquatic environments have tested positive for elevated levels of
toxins A toxin is a naturally occurring poison produced by metabolic activities of living cells or organisms. They occur especially as proteins, often conjugated. The term was first used by organic chemist Ludwig Brieger (1849–1919), derived ...
, such as mercury,
arsenic Arsenic is a chemical element; it has Symbol (chemistry), symbol As and atomic number 33. It is a metalloid and one of the pnictogens, and therefore shares many properties with its group 15 neighbors phosphorus and antimony. Arsenic is not ...
,
chromium Chromium is a chemical element; it has Symbol (chemistry), symbol Cr and atomic number 24. It is the first element in Group 6 element, group 6. It is a steely-grey, Luster (mineralogy), lustrous, hard, and brittle transition metal. Chromium ...
, and
copper Copper is a chemical element; it has symbol Cu (from Latin ) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkish-orang ...
, there are posted signs with warnings about the consumption of fish caught in those areas. Concentration of mercury biomagnifies as it passes up the food chain from organisms on lower trophic levels to apex predators. It
bioaccumulate Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance faster than it can be lost or eliminated by catabolism and excretion. Th ...
s in the tissues of larger, long-lived predatory fish. When compared to smaller, short-lived fish, bowfin tend to concentrate mercury at higher levels thereby making them less safe for human consumption.


See also

*
Snakehead (fish) The snakeheads are members of the freshwater perciform fish family Channidae, native to parts of Africa and Asia. These elongated, predatory fish are distinguished by their long dorsal fins, large mouths, and shiny teeth. They breathe air w ...


References


Further reading

* * * * *


External links


MN DNR

Bowfin Anglers' Group

FishBase info for Amiidae


{{Taxonbar, from=Q491934 Amia (fish)
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
Fish of the Great Lakes Freshwater fish of the Southeastern United States Articles containing video clips Fish described in 1766