The anamniotes are an
informal group of
craniates comprising all
fish
A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
and
amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, which lay their
egg
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
s in
aquatic environment
An aquatic ecosystem is an ecosystem found in and around a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms—aquatic life—that are dependent on each other and on their environm ...
s. They are distinguished from the
amniote
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s (
reptile
Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s,
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s and
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s), which can reproduce on
dry land
Land, also known as dry land, ground, or earth, is the solid terrestrial surface of Earth not submerged by the ocean or another body of water. It makes up 29.2% of Earth's surface and includes all continents and islands. Earth's land surface ...
either by laying
shell
Shell may refer to:
Architecture and design
* Shell (structure), a thin structure
** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses
Science Biology
* Seashell, a hard outer layer of a marine ani ...
ed eggs or by carrying
fertilized egg
A zygote (; , ) is a eukaryotic cell formed by a fertilization event between two gametes.
The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism.
The sexual ...
s
within the female. Older sources, particularly before the 20th century, may refer to anamniotes as "lower vertebrates" and amniotes as "higher vertebrates", based on the antiquated idea of the
evolutionary great chain of being.
The name "anamniote" is a
back-formation
Back-formation is the process or result of creating a neologism, new word via Morphology (linguistics), morphology, typically by removing or substituting actual or supposed affixes from a lexical item, in a way that expands the number of lexemes ...
word created by adding the prefix ''
an-'' to the word ''amniote'', which in turn refers to the
amnion
The amnion (: amnions or amnia) is a membrane that closely covers human and various other embryos when they first form. It fills with amniotic fluid, which causes the amnion to expand and become the amniotic sac that provides a protective envir ...
, an
extraembryonic membrane The extraembryonic membranes are four membranes which assist in the development of an animal's embryo. Such membranes occur in a range of animals from humans to insects. They originate from the zygote, but are not considered part of the embryo. The ...
present during the amniotes'
embryonic development
In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm, sperm cell (spermat ...
which serves as a biochemical barrier that shields the
embryo
An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
from environmental fluctuations by regulating the
oxygen
Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
,
carbon dioxide
Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
and
metabolic waste
Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds ...
exchanges and secreting
a cushioning fluid. As the name suggests, anamniote embryos lack an amnion during embryonic development, and therefore rely on the presence of external water to provide oxygen and help dilute and excrete waste products (particularly
ammonia
Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
) via
diffusion
Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
in order for the embryo to complete development without being intoxicated by their own metabolites.
This means anamniotes are almost always dependent on an aqueous (or at least very moist) environment for
reproduction
Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
and are thus restricted to
spawning
Spawn is the Egg cell, eggs and Spermatozoa, sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of freely releasing eggs and sperm into a body of water (fresh or marine); the physical act is ...
in or near
water bodies. They are also highly sensitive to chemical and temperature variation in the surrounding water, and are also more vulnerable to
egg predation and
parasitism
Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
.
During their
life cycle, all anamniote
class
Class, Classes, or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used d ...
es pass through a completely aquatic egg stage, as well as an aquatic
larva
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l stage during which all
hatchlings are
gill
A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...
-dependent and morphologically resemble tiny
finless fish (known as a
fry or a
tadpole
A tadpole or polliwog (also spelled pollywog) is the Larva, larval stage in the biological life cycle of an amphibian. Most tadpoles are fully Aquatic animal, aquatic, though some species of amphibians have tadpoles that are terrestrial animal, ...
for fish and amphibians, respectively), before
metamorphosizing into
juvenile and
adult
An adult is an animal that has reached full growth. The biological definition of the word means an animal reaching sexual maturity and thus capable of reproduction. In the human context, the term ''adult'' has meanings associated with social an ...
forms (which might be
aquatic,
semiaquatic
In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
or even
terrestrial), thus indicating their
physiological
Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
homology.
Anamniote traits

The group is characterized by retaining the primitive vertebrate condition in several traits:
* Absence of an
amnion
The amnion (: amnions or amnia) is a membrane that closely covers human and various other embryos when they first form. It fills with amniotic fluid, which causes the amnion to expand and become the amniotic sac that provides a protective envir ...
* Absence or rudimentary condition of the
allantois
* Permeable skin allowing diffusion of water and gases directly through the skin.
* Presence at some period of life of
gills.
History of discovery
The features unifying the anamniotes was first noted by
Thomas Henry Huxley in 1863, who coined the phrase ''Ichtioid'' or ''Ichthyopsida'' ("fish-face") for the group. It is a
taxonomic classification just below the level of
Vertebrata
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 Verte ...
, though Huxley presented the Ichthyopsida as an informal unit and never ventured to forward a
Linnaean rank for the group. The term Ichthyopsida means ''fish-face'' or ''fish-like'' as opposed to the
Sauropsida
Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
or ''lizard-face'' animals (
reptiles
Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
and
birds
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
) and the
mammals
A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
.
[Encyclopædia Britannica, 9th ed. (1878)]
original text
/ref>
The group representing an evolutionary grade
A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit.
Phylogenetics
The concept of evolutionary grades ...
rather than a 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 ...
, the term anamniote is now used as an informal way of denoting the physical property of the group, rather than as a systematic unit.
References
{{Taxonbar, from=Q485501
Zoology
Paraphyletic groups