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''Amphiuma'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of aquatic
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s from the United States, the only
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
genus within the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Amphiumidae . They are colloquially known as amphiumas. They are also known to fishermen as "
conger eel ''Conger'' ( ) is a genus of marine congrid eels. It includes some of the largest types of eels, ranging up to 2 m (6 ft) or more in length, in the case of the European conger. Large congers have often been observed by divers during t ...
s" or "Congo snakes", which are zoologically incorrect designations or
misnomer A misnomer is a name that is incorrectly or unsuitably applied. Misnomers often arise because something was named long before its correct nature was known, or because an earlier form of something has been replaced by a later form to which the name ...
s, since amphiumas are actually
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s (and thus
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s), and not
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 ...
, nor
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s and are not from Congo. ''Amphiuma'' exhibits one of the largest complements of DNA in the living world, around 25 times more than a human.


Taxonomy

Numerous phylogenetic studies have indicated that amphiumas form a clade with the families
Rhyacotritonidae The torrent salamanders or Cascade salamanders are a family of salamanders (Rhyacotritonidae) with only one genus, ''Rhyacotriton''. The torrent salamanders are endemic to the United States in the Pacific Northwest (including northwestern Calif ...
(torrent salamanders) and
Plethodontidae Plethodontidae, or lungless salamanders, are a family of salamanders. Most species are native to the Western Hemisphere, from British Columbia to Brazil, although a few species are found in Sardinia, Europe south of the Alps, and South Korea. In ...
(lungless salamanders), with an especially close relationship to Plethodontidae. Despite this possible relationship, the two families must have still diverged very early on. The genus ''Proamphiuma'' from the Cretaceous is the earliest known member of the family, and closely resembles the modern species aside from less elaborate vertebral structures.


Description

Amphiumas have an elongated body, generally grey-black in color. They do have legs, but they are merely vestigial and very small. While amphiumas can be up to long, their legs measure only up to about . It is because of this that they are often mistaken for
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s or snakes. They also lack eyelids and a tongue. Amphiumas also have a lateral line visible on the sides of their bodies, which is capable of detecting movement and is used in aid of hunting. Female amphiumas lay their
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
in wet mud, and then remain coiled around them for about five months, until they hatch. The larvae have external gills, but after about four months these external gills disappear and the lungs begin to work. One pair of
gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
s, with fully functioning internal gills, is retained and never disappears, so the
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
remains incomplete.


Species

There are three extant amphiuma species, distinguished by the number of toes:


Extant


Extinct

* †''
Amphiuma jepseni ''Amphiuma'' is a genus of aquatic salamanders from the United States, the only extant genus within the family Amphiumidae . They are colloquially known as amphiumas. They are also known to fishermen as "conger eels" or "Congo snakes", which are ...
'' (
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
of
Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the south ...
) * †''
Amphiuma antica ''Amphiuma'' is a genus of aquatic salamanders from the United States, the only extant genus within the family Amphiumidae . They are colloquially known as amphiumas. They are also known to fishermen as "conger eels" or "Congo snakes", which are ...
'' (
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
)


Distribution

Amphiumas inhabit the southeastern part of the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
. They share much of the same distribution with the
sirens Siren or sirens may refer to: Common meanings * Siren (alarm), a loud acoustic alarm used to alert people to emergencies * Siren (mythology), an enchanting but dangerous monster in Greek mythology Places * Siren (town), Wisconsin * Siren, Wisco ...
, although they are not closely related. In the past, amphiumas had a wider geographic range throughout North America, ranging all the way north to
Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the south ...
.


Behavior

During the day, amphiumas hide in vegetation, and at night they become active hunters. Their prey includes
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,
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s,
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 ...
,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and even other amphiumas. Hunting and eating habits have been observed to be very similar to that of the
axolotl The axolotl (; from nci, āxōlōtl ), ''Ambystoma mexicanum'', is a paedomorphic salamander closely related to the tiger salamander. Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. Instea ...
, including the sucking in of food by their stomachs with vacuum force. If provoked, they can become aggressive. They can be found in most wetlands in the coastal plain of the southeastern U.S., even ones which periodically dry out, as they are able to estivate in the moist mud below drained marshland and other
ephemeral Ephemerality (from the Greek word , meaning 'lasting only one day') is the concept of things being transitory, existing only briefly. Academically, the term ephemeral constitutionally describes a diverse assortment of things and experiences, fr ...
wetlands. Amphiumas are rarely encountered on land.


Food habits

The amphiuma's predatory behaviors and food selection are very calculated and variable depending on abundance of food. In addition to eating
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,
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s,
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 ...
,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, and other amphiuma, amphiuma have been found to eat
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s,
vegetable Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems, ...
s,
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroon ...
s,
mollusca Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
, and
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
. Amphiuma seem to have a preference for eating
crawfish Crayfish are freshwater crustaceans belonging to the clade Astacidea, which also contains lobsters. In some locations, they are also known as crawfish, craydids, crawdaddies, crawdads, freshwater lobsters, mountain lobsters, rock lobsters, mu ...
. It has been documented that amphiuma will pass on smaller crawfish in order to consume larger ones. It is suggested that this limits wasting energy in pursuing prey with less caloric density. In captivity, the predatory behavior amphiuma display depends on the presence or lack of food. Amphiuma will remain inactive when food is absent, and will become more active once food has been introduced into their habitat. This shows that the amphiuma, although ancestral to many
amphibia Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...
, has developed a deductive approach to its predation.


Anatomy


Jaw muscles

Amphiuma are primarily carnivorous amphibians that consume crayfish, insects, and other small invertebrates. Similar to many
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s, the amphiuma has two distinct forms of suction feeding procedures: stationary and strike. Stationary suction feeding involve little to no movement where it opens the mouth with buccal expansion but no forward movement of the body. strike suction is rapid motion where the mouth opens and buccal expansion occurs synchronously leading to a quick attack. These two feeding habits give the amphiuma the ability to have a larger variety of food (live or dead). amphiuma's ability to displace its jaw to feed means they can consume a large variety of organisms as well. But amphiuma's narrow jaw makes it harder for them to fully consume large prey such as crayfish or mice. In these cases, they will use one of the forms of suction feeding and then rip the prey into pieces until fully consumed. Small prey will be pulled completely into the mouth before being eaten. Structure of the teeth within the jaw tend to be arched caudal on the head. The muscles of the jaw give them the ability to grab and hold on to prey as well as create a negative pressure to suck in the prey and displace the jaw. Muscles of note in the amphiuma include: the levator mandibulae anterior and levator mandibulae externus, which elevate the lower jaw of the amphiuma while the depressor mandibulae depresses the lower jaw. The intermandubularis works by tensing the mouth floor. The branchiohyoideus and geniohyoideus draws the hyoid arch which causes the suction and displacement.


Lungs

Amphiuma possess relatively ancestral forms of lungs compared to some of the other groups of
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s that live terrestrially today. Their lungs are long organs, extending over half of the body length, with dense capillary networks and large surface area that suggest the utilization of the entire lung for respiration while the animal is in water or on land. Although it is common for amphibia to respire out of their skin, also known as
cutaneous respiration Cutaneous respiration, or cutaneous gas exchange (sometimes called, skin breathing), is a form of respiration in which gas exchange occurs across the skin or outer integument of an organism rather than gills or lungs. Cutaneous respiration may be th ...
, it was found that amphiuma primarily respire through their lungs, despite their aquatic lifestyle. This is suggested by the high lung to respiratory capillary density compared to the relatively low skin to respiratory capillary density.


Respiration

Pressure gradients for respiration occur in two different locations, the buccal cavity/nares (mouth and nostril) region, and in the lungs of the amphiuma. The first system for respiration occurs in the buccal cavity/nares through a two-cycle pressure-induced buccal cavity/nares process. In the first system, the amphiuma performs one full cycle of body expansion and compression in order to inhale and another full cycle to exhale, a unique process that utilizes both the
buccal cavity The buccal space (also termed the buccinator space) is a fascial spaces of the head and neck, 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 ...
and the nares (openings of nostrils). The buccal cavity creates pressure that aids in driving the cycles of expansion and compression required for respiration, although it was found that the buccal pressure gradient alone was not enough to drive respiration in the ''
Amphiuma tridactylum ''Amphiuma tridactylum'', the three-toed amphiuma, is a species of aquatic salamander native to the Southeastern United States. Description The three-toed amphiuma looks rather eel-like, with an elongate, dark gray-black, or brown colored b ...
''. Rather, the buccal cavity allows for small pressure changes that are thought to have an
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
purpose. This buccal cavity/nares component to the amphiuma respiratory process supplements the contribution performed by the lung. It is the pressure control performed in the lungs that drive the inhalation and exhalation forces through the flexing of
smooth muscle Smooth muscle is an involuntary non-striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit mus ...
in the lung. In order to exhale, amphiuma push air from their lungs into their buccal cavity, distending the cavity, before releasing the air. Without inhaling, amphiuma repeat the process, exhaling a second volume of air that allows them to completely empty their lungs. Only after both exhales can they then inhale, using a negative pressure gradient made by the smooth muscles in their lungs to take in air.


Sexual dimorphism

Amphiuma demonstrate
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
in relation to the size of their bodies and the size of their heads. Generally, males have been found to possess larger bodies and longer heads compared to the female sex, which normally is indicative of male-male combat observed within the population. There has been, however, no other physical indicating factors for male-male combat as in other species of amphibians, such as horns or spines. Some populations do not show these sexual dimorphic traits, and in certain locations female and male bodies do not exhibit any traits with significant differences. Amphiumas may be sexed as male or female based on the pigmentation of the cloacal opening. Males exhibit white or pink coloration while females exhibit dark pigmentation. Occasionally, males may demonstrate partial pigmentation, but never have full dark coloration like that of females.


References


External links


Tree of Life: Amphiumidaeamphiumas.org
{{Taxonbar, from=Q300923 Salamandroidea Extant Pleistocene first appearances Amphibian genera Endemic fauna of the United States Taxa named by Alexander Garden (naturalist)