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Marine invertebrates are the
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s that live in
marine habitat Marine habitats are habitats that support marine life. Marine life depends in some way on the saltwater that is in the sea (the term ''marine'' comes from the Latin ''mare'', meaning sea or ocean). A habitat is an ecological or environmental ...
s. Invertebrate is a blanket term that includes all
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s apart from the
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
members of the
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
phylum.
Invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s lack a
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
, and some have evolved a
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 ** Thin-shell structure Science Biology * Seashell, a hard o ...
or a hard
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
. As on land and in the air, marine invertebrates have a large variety of
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
s, and have been categorised into over 30
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
. They make up most of the macroscopic life in the oceans.


Evolution

The earliest
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s were marine
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s, that is,
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s came later. Animals are
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
s, and are distinguished from plants, algae, and fungi by lacking
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
s. Marine invertebrates are animals that inhabit a
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military ...
environment apart from the vertebrate members of the
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
phylum; invertebrates lack a
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
. Some have evolved a
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 ** Thin-shell structure Science Biology * Seashell, a hard o ...
or a hard
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
. The earliest animals may belong to the genus ''
Dickinsonia ''Dickinsonia'' is an extinct genus of basal animal that lived during the late Ediacaran period in what is now Australia, China, Russia and Ukraine. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its a ...
'', 571 million to 539 million years ago. Individual ''Dickinsonia'' typically resemble a bilaterally symmetrical ribbed oval. They kept growing until they were covered with sediment or otherwise killed, and spent most of their lives with their bodies firmly anchored to the sediment. Their taxonomic affinities are presently unknown, but their mode of growth is consistent with a
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
n affinity. Apart from '' Dickinsonia'', the earliest widely accepted animal fossils are the rather modern-looking
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns (the group that includes
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbre ...
,
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, ...
s and '' Hydra''), possibly from around The
Ediacara biota The Ediacaran (; formerly Vendian) biota is a taxonomic period classification that consists of all life forms that were present on Earth during the Ediacaran Period (). These were composed of enigmatic tubular and frond-shaped, mostly sessil ...
, which flourished for the last 40 million years before the start of the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
, were the first animals more than a very few centimetres long. Like '' Dickinsonia'', many were flat with a "quilted" appearance, and seemed so strange that there was a proposal to classify them as a separate kingdom,
Vendozoa Vendobionts or Vendozoans (Vendobionta) are a proposed very high-level, extinct clade of benthic organisms that made up of the majority of the organisms that were part of the Ediacaran biota. It is a hypothetical group and at the same time, it ...
. Others, however, have been interpreted as early
mollusc 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 est ...
s (''
Kimberella ''Kimberella'' is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with t ...
''),
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s ('' Arkarua''), and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s (''
Spriggina ''Spriggina'' is a genus of early bilaterian animals whose relationship to living animals is unclear. Fossils of ''Spriggina'' are known from the late Ediacaran period in what is now South Australia. ''Spriggina floundersi'' is the official fo ...
'', '' Parvancorina''). There is still debate about the classification of these specimens, mainly because the diagnostic features which allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the Ediacarans. However, there seems little doubt that ''Kimberella'' was at least a
triploblastic Triploblasty is a condition of the gastrula in which there are three primary germ layers: the ectoderm, mesoderm, and endoderm. Germ cells are set aside in the embryo at the blastula stage, which are incorporated into the gonads during organo ...
bilaterian animal, in other words, an animal significantly more complex than the cnidarians. The
small shelly fauna The small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) are mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end of the Early Cambri ...
are a very mixed collection of fossils found between the Late Ediacaran and
Middle Cambrian Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ( ...
periods. The earliest, '' Cloudina'', shows signs of successful defense against predation and may indicate the start of an
evolutionary arms race In evolutionary biology, an evolutionary arms race is an ongoing struggle between competing sets of co-evolving genes, phenotypic and behavioral traits that develop escalating adaptations and counter-adaptations against each other, resembling an ...
. Some tiny Early Cambrian shells almost certainly belonged to molluscs, while the owners of some "armor plates," ''
Halkieria The halkieriids are a group of fossil organisms from the Lower to Middle Cambrian. Their eponymous genus is ''Halkieria'' , which has been found on almost every continent in Lower to Mid Cambrian deposits, forming a large component of the sma ...
'' and ''
Microdictyon ''Microdictyon'' is an extinct armoured worm-like animal coated with net-like scleritic plates, known from the Early Cambrian Maotianshan shale of Yunnan China and other parts of the world. ''Microdictyon'' is part of the ill-defined taxon &ndas ...
'', were eventually identified when more complete specimens were found in Cambrian
lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n that preserved soft-bodied animals. File:DickinsoniaCostata.jpg, ''
Dickinsonia ''Dickinsonia'' is an extinct genus of basal animal that lived during the late Ediacaran period in what is now Australia, China, Russia and Ukraine. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its a ...
'' may be the earliest animal. They appear in the fossil record 571 million to 541 million years ago. File:Kimberella NT.jpg, ''
Kimberella ''Kimberella'' is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with t ...
'', an early mollusc important for understanding the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
. Invertebrates are grouped into different
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
(
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
s). File:20191108 Opabinia regalis.png, '' Opabinia'', an extinct
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. ...
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
appeared in the
Middle Cambrian Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ( ...
In the 1970s there was already a debate about whether the emergence of the modern phyla was "explosive" or gradual but hidden by the shortage of
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of th ...
animal fossils. A re-analysis of fossils from the
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fo ...
lagerstätte increased interest in the issue when it revealed animals, such as '' Opabinia'', which did not fit into any known
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
. At the time these were interpreted as evidence that the modern phyla had evolved very rapidly in the Cambrian explosion and that the Burgess Shale's "weird wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution. Later discoveries of similar animals and the development of new theoretical approaches led to the conclusion that many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups—for example that ''Opabinia'' was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it may have been closely related to the modern
tardigrade Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserb� ...
s. Nevertheless, there is still much debate about whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it appears unique in the history of animals.


Classification

Invertebrates are grouped into different
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
. Informally phyla can be thought of as a way of grouping organisms according to their
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
. A body plan refers to a blueprint which describes the shape or morphology of an organism, such as its
symmetry Symmetry (from grc, συμμετρία "agreement in dimensions, due proportion, arrangement") in everyday language refers to a sense of harmonious and beautiful proportion and balance. In mathematics, "symmetry" has a more precise definiti ...
, segmentation and the disposition of its
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ante ...
s. The idea of body plans originated with
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, which were grouped into one phylum. But the vertebrate body plan is only one of many, and invertebrates consist of many phyla or body plans. The history of the discovery of body plans can be seen as a movement from a worldview centred on vertebrates, to seeing the vertebrates as one body plan among many. Among the pioneering
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
s, Linnaeus identified two body plans outside the vertebrates; Cuvier identified three; and Haeckel had four, as well as the Protista with eight more, for a total of twelve. For comparison, the number of phyla recognised by modern zoologists has risen to 35. Historically body plans were thought of as having evolved in rapidly during the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
, but a more nuanced understanding of animal evolution suggests a gradual development of body plans throughout the early
Palaeozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and '' ...
and beyond. More generally a phylum can be defined in two ways: as described above, as a group of organisms with a certain degree of morphological or developmental similarity (the
phenetic In biology, phenetics ( el, phainein – to appear) , also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary re ...
definition), or a group of organisms with a certain degree of evolutionary relatedness (the
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
definition). As on land and in the air, invertebrates make up a great majority of all macroscopic life, as the vertebrates makes up a subphylum of one of over 30 known animal phyla, making the term almost meaningless for taxonomic purpose. Invertebrate sea life includes the following groups, some of which are phyla: *
Acoela Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order o ...
, among the most primitive
bilateral animals The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
; * Annelida, (
polychaete Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made ...
s and sea
leech Leeches are segmented parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
es); * Brachiopoda, marine animals that have hard "valves" (shells) on the upper and lower surfaces ; *
Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
, also known as moss animals or sea mats; *
Chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and ca ...
, commonly known as arrow worms, are a phylum of predatory marine worms that are a major component of plankton; *
Cephalochordata A cephalochordate (from Greek: κεφαλή ''kephalé'', "head" and χορδή ''khordé'', "chord") is an animal in the chordate subphylum, Cephalochordata. They are commonly called lancelets. Cephalochordates possess 5 synapomorphies, or prima ...
represented in the modern oceans by the
lancelets The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
(also known as Amphioxus); *
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
, such as
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbre ...
,
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, ...
s, and
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
s; *
Crustacea 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 c ...
, including
lobster Lobsters are a family (Nephropidae, synonym Homaridae) of marine crustaceans. They have long bodies with muscular tails and live in crevices or burrows on the sea floor. Three of their five pairs of legs have claws, including the first pair, ...
s,
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all th ...
s,
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are ref ...
,
crayfish 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, m ...
,
barnacle A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosiv ...
s,
hermit crab Hermit crabs are anomuran decapod crustaceans of the superfamily Paguroidea that have adapted to occupy empty scavenged mollusc shells to protect their fragile exoskeletons. There are over 800 species of hermit crab, most of which possess an as ...
s,
mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length, ...
s, and
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have ...
s; *
Ctenophora Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
, also known as comb jellies, the largest animals that swim by means of cilia; *
Echinodermata An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the s ...
, including
sea star Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish a ...
s,
brittle star Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
s,
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) o ...
s,
sand dollar Sand dollars (also known as a sea cookie or snapper biscuit in New Zealand, or pansy shell in South Africa) are species of flat, burrowing sea urchins belonging to the order Clypeasteroida. Some species within the order, not quite as flat, are k ...
s,
sea cucumber Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuri ...
s,
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
s, and sea daisies; *
Echiura The Echiura, or spoon worms, are a small group of marine animals. Once treated as a separate phylum, they are now considered to belong to Annelida. Annelids typically have their bodies divided into segments, but echiurans have secondarily los ...
, also known as spoon worms; * Gnathostomulids, slender to thread-like worms, with a transparent body that inhabit sand and mud beneath shallow coastal waters; * Gastrotricha, often called hairy backs, found mostly interstitially in between sediment particles; * Hemichordata, includes
acorn worms The acorn worms or Enteropneusta are a hemichordate class of invertebrates consisting of one order of the same name. The closest non-hemichordate relatives of the Enteropneusta are the echinoderms. There are 111 known species of acorn worm in the ...
, solitary worm-shaped organisms; * Kamptozoa, goblet-shaped sessile aquatic animals, with relatively long stalks and a "crown" of solid tentacles, also called Entoprocta; *
Kinorhyncha Kinorhyncha ( grc, κινέω, kīnéō, I move, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are also called mud dragons. Modern species are or less, ...
, segmented, limbless animals, widespread in mud or sand at all depths, also called mud dragons; * Loricifera, very small to microscopic
marine sediment Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles have their origins in soil and rocks and have been transported from the land to the sea, mai ...
-dwelling animals only discovered in 1983; *
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 es ...
, including
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater environ ...
,
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fittin ...
,
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, ...
,
whelk Whelk (also known as scungilli) is a common name applied to various kinds of sea snail. Although a number of whelks are relatively large and are in the family Buccinidae (the true whelks), the word ''whelk'' is also applied to some other marin ...
s, ''
Nautilus The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in ...
'',
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
,
nudibranch Nudibranchs () are a group of soft-bodied marine gastropod molluscs which shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to match, ...
s,
scallop Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
s,
sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the a ...
s,
Aplacophora Aplacophora is a presumably paraphyletic taxon. This is a class of small, deep-water, exclusively benthic, marine molluscs found in all oceans of the world. MolluscaBase eds. (2021). MolluscaBase. Aplacophora. Accessed through: World Register of ...
, Caudofoveata,
Monoplacophora Monoplacophora , meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell inhabiting deep sea environments . Extant representatives were not recognized as such until 1952; previously they were known only from ...
, Polyplacophora, and Scaphopoda; * Myzostomida, a taxonomic group of small marine worms which are parasitic on crinoids or "sea lilies"; *
Nemertinea Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many h ...
, also known as "ribbon worms" or "proboscis worms"; * Orthonectida, a small phylum of poorly known parasites of marine invertebrates that are among the simplest of multi-cellular organisms; * Phoronida, a phylum of marine animals that filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to support and protect their soft bodies; *
Placozoa The Placozoa are a basal form of marine free-living (non-parasitic) multicellular organism. They are the simplest in structure of all animals. Three genera have been found: the classical ''Trichoplax adhaerens'', '' Hoilungia hongkongensis'', a ...
, small, flattened, multicellular animals around 1 millimetre across and the simplest in structure. They have no regular outline, although the lower surface is somewhat concave, and the upper surface is always flattened; *
Porifera Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
(sponges), multicellular organisms that have bodies full of pores and channels allowing water to circulate through them; * Priapulida, or penis worms, are a phylum of marine worms that live marine mud. They are named for their extensible spiny proboscis, which, in some species, may have a shape like that of a human penis; *
Pycnogonida Sea spiders are marine arthropods of the order Pantopoda ( ‘all feet’), belonging to the class Pycnogonida, hence they are also called pycnogonids (; named after ''Pycnogonum'', the type genus; with the suffix '). They are cosmopolitan, foun ...
, also called sea spiders, are unrelated to spiders, or even to arachnids which they resemble; *
Sipunculida The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a class containing about 162 species of unsegmented marine annelid worms. The name ''Sipuncula'' is from the genus name '' Sipunculus'', and comes from the Latin ...
, also called peanut worms, is a group containing 144–320 species (estimates vary) of bilaterally symmetrical, unsegmented marine worms; * Tunicata, also known as sea squirts or sea pork, are filter feeders attached to rocks or similarly suitable surfaces on the ocean floor; *Some
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegmen ...
s of the classes
Turbellaria The Turbellaria are one of the traditional sub-divisions of the phylum Platyhelminthes (flatworms), and include all the sub-groups that are not exclusively parasitic. There are about 4,500 species, which range from to large freshwater forms mo ...
and
Monogenea Monogeneans are a group of ectoparasitic flatworms commonly found on the skin, gills, or fins of fish. They have a direct lifecycle and do not require an intermediate host. Adults are hermaphrodites, meaning they have both male and female repr ...
; * Xenoturbella, a genus of bilaterian animals that contains only two marine worm-like species; *
Xiphosura Xiphosura () is an order of arthropods related to arachnids. They are more commonly known as horseshoe crabs (a name applied more specifically to the only extant family, Limulidae). They first appeared in the Hirnantian ( Late Ordovician). Curr ...
, includes a large number of extinct lineages and only four recent species in the family Limulidae, which include the
horseshoe crab Horseshoe crabs are marine and brackish water arthropods of the family Limulidae and the only living members of the order Xiphosura. Despite their name, they are not true crabs or crustaceans: they are chelicerates, most closely related to ar ...
s.
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s total about 1,113,000 described extant species,
mollusc 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 est ...
s about 85,000 and
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
s about 52,000.


Marine sponges

Sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...
s are animals of the
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
Porifera (Modern Latin for ''bearing pores'' Porifera (n.)
''Online Etymology Dictionary''. Retrieved 18 August 2016.
). They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
sandwiched between two thin layers of
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
s. They have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
s. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. Sponges are similar to other animals in that they are
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
,
heterotrophic A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
, lack
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
s and produce
sperm cell Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
s. Unlike other animals, they lack true tissues and organs, and have no
body symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...
. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where it deposits nutrients, and leaves through a hole called the
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
. Many sponges have internal skeletons of
spongin Spongin, a modified type of collagen protein, forms the fibrous skeleton of most organisms among the phylum Porifera, the sponges. It is secreted by sponge cells known as spongocytes. Spongin gives a sponge its flexibility. True spongin is found ...
and/or spicules of calcium carbonate or
silicon dioxide Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
. All sponges are sessile aquatic animals. Although there are freshwater species, the great majority are marine (salt water) species, ranging from tidal zones to depths exceeding . While most of the approximately 5,000–10,000 known species feed on
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
and other food particles in the water, some host
photosynthesizing Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
micro-organisms as
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s and these alliances often produce more food and oxygen than they consume. A few species of sponge that live in food-poor environments have become
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s that prey mainly on small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s. File:Sponges in Caribbean Sea, Cayman Islands.jpg, Sponge biodiversity. There are four sponge species in this photo. File:Callyspongia vaginalis (Branching Vase Sponge - pink variation).jpg,
Branching vase sponge ''Callyspongia'' (''Cladochalina'') ''aculeata'', commonly known as the branching vase sponge is a species of '' Porifora'', meaning sea sponge, in the family Callyspongiidae. Poriferans are typically characterized by ostia, pores that filter out ...
File:Euplectella aspergillum (cropped).jpg, Venus' flower basket at a depth of 2572 meters File:Barrel sponge (Xestospongia testudinaria).jpg, Barrel sponge File:Aplysina archeri (Stove-pipe Sponge-pink variation).jpg, Stove-pipe sponge
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
mistakenly identified sponges as plants in the order
Algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
. For a long time thereafter sponges were assigned to a separate subkingdom,
Parazoa Parazoa (Parazoa, gr. Παρα-, para, "next to", and ζωα, zoa, "animal") are a taxon with sub-kingdom category that is located at the base of the phylogenetic tree of the animal kingdom in opposition to the sub-kingdom Eumetazoa; they group ...
(meaning ''beside the animals''). They are now classified as a
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
from which the higher animals have evolved.


Marine cnidarians

Cnidarians Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that the ...
(Greek for ''nettle'') are distinguished by the presence of stinging cells, specialized cells that they use mainly for capturing prey. Cnidarians include
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
s,
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, ...
s,
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbre ...
and
hydrozoans Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specialize ...
. They form a
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
containing over 10,000
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
of
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s found exclusively in aquatic (mainly marine) environments. Their bodies consist of
mesoglea Mesoglea refers to the extracellular matrix found in cnidarians like coral or jellyfish that functions as a hydrostatic skeleton. It is related to but distinct from mesohyl, which generally refers to extracellular material found in sponges. Desc ...
, a non-living jelly-like substance, sandwiched between two layers of
epithelium Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...
that are mostly one cell thick. They have two basic body forms: swimming
medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
e and sessile polyps, both of which are radially symmetrical with mouths surrounded by tentacles that bear cnidocytes. Both forms have a single orifice and body cavity that are used for digestion and
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
. Fossil cnidarians have been found in rocks formed about . Fossils of cnidarians that do not build mineralized structures are rare. Scientists currently think cnidarians,
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), an ...
s and
bilaterian The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...
s are more closely related to calcareous sponges than these are to other
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...
s, and that
anthozoan Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of t ...
s are the evolutionary "aunts" or "sisters" of other cnidarians, and the most closely related to bilaterians.
Cnidarians Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that the ...
are the simplest animals in which the cells are organised into tissues. The
starlet sea anemone The starlet sea anemone (''Nematostella vectensis'') is a species of small sea anemone in the family Edwardsiidae native to the east coast of the United States, with introduced populations along the coast of southeast England and the west coast ...
is used as a
model organism A model organism (often shortened to model) 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 workin ...
in research. It is easy to care for in the laboratory and a
protocol Protocol may refer to: Sociology and politics * Protocol (politics), a formal agreement between nation states * Protocol (diplomacy), the etiquette of diplomacy and affairs of state * Etiquette, a code of personal behavior Science and technology ...
has been developed which can yield large numbers of embryos on a daily basis. There is a remarkable degree of similarity in the gene sequence conservation and complexity between the sea anemone and vertebrates. In particular, genes concerned in the formation of the head in vertebrates are also present in the anemone. File:Sea anemone in tidepools.jpg, Sea anemones are common in tidepools File:Striped colonial anemone.jpg, Their
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s sting and paralyse small fish File:Coral detail.jpg, Close up of polyps on the surface of a
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
, waving their tentacles. File:Maldives small island.jpg, If an island sinks below the sea, coral growth can keep pace with the rising water and form an
atoll An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon partially or completely. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical oceans and seas where corals can gro ...
File:Portuguese Man-O-War (Physalia physalis).jpg,
Portuguese man o' war The Portuguese man o' war (''Physalia physalis''), also known as the man-of-war, is a marine hydrozoan found in the Atlantic Ocean and the Indian Ocean. It is considered to be the same species as the Pacific man o' war or blue bottle, which is ...
File:Porpita porpita.jpg, ''
Porpita porpita ''Porpita porpita'', or blue button, is a marine organism consisting of a colony of hydroids found in the warmer, tropical and sub-tropical waters of the Pacific, Atlantic, and Indian oceans, as well as the Mediterranean Sea and eastern Arabian ...
'' File:Largelionsmanejellyfish.jpg, Lion's mane jellyfish, largest known jellyfish File:Sea walnut, Boston Aquarium (cropped).jpg, The
bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some b ...
sea walnut ''Mnemiopsis leidyi'', the warty comb jelly or sea walnut, is a species of tentaculate ctenophore (comb jelly). It is native to western Atlantic coastal waters, but has become established as an invasive species in European and western Asian regi ...
has a transient anus which forms only when it needs to defecate File:Turritopsis dohrnii (cropped).jpg, ''
Turritopsis dohrnii ''Turritopsis dohrnii'', also known as the immortal jellyfish, is a species of small, biologically immortal jellyfish found worldwide in temperate to tropic waters. It is one of the few known cases of animals capable of reverting completely to ...
'' achieves
biological immortality Biological immortality (sometimes referred to as bio-indefinite mortality) is a state in which the rate of mortality from senescence is stable or decreasing, thus decoupling it from chronological age. Various unicellular and multicellular species, ...
by transferring its cells back to childhood


Marine worms

Worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in size from microscopic to over in length for marine polychaete wor ...
s (Old English for ''serpent'') typically have long cylindrical tube-like bodies and no
limb Limb may refer to: Science and technology * Limb (anatomy), an appendage of a human or animal *Limb, a large or main branch of a tree *Limb, in astronomy, the curved edge of the apparent disk of a celestial body, e.g. lunar limb *Limb, in botany, ...
s.
Marine worm Any worm that lives in a marine environment is considered a marine worm. Marine worms are found in several different phyla, including the Platyhelminthes, Nematoda, Annelida (segmented worms), Chaetognatha, Hemichordata, and Phoronida. For a ...
s vary in size from microscopic to over in length for some marine polychaete worms (
bristle worms Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are ma ...
) and up to for the marine nemertean worm (
bootlace worm The bootlace worm (''Lineus longissimus'') is a species of ribbon worm and one of the longest known animals, with specimens up to long being reported, although this has not been confirmed. Its mucus is highly toxic. Taxonomy The bootlace worm ...
).
Mark Carwardine Mark Carwardine ( IPA: /kɑːwɑːdiːn/; born 9 March 1959) is a British zoologist who achieved widespread recognition with his 20-year conservation project – '' Last Chance to See'' – which involved round-the-world expeditions with Dougla ...
(1995) ''The Guinness Book of Animal Records''. Guinness Publishing. p. 232.
Some marine worms occupy a small variety of
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
niches, living inside the bodies of other animals, while others live more freely in the marine environment or by burrowing underground. Different groups of marine worms are related only distantly, so they are found in several different
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
such as the Annelida (segmented worms),
Chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and ca ...
(arrow worms), Hemichordata, and Phoronida (horseshoe worms). Many of these worms have specialized tentacles used for exchanging oxygen and carbon dioxide and also may be used for reproduction. Some marine worms are
tube worm A tubeworm is any worm-like sessile invertebrate that anchors its tail to an underwater surface and secretes around its body a mineral tube, into which it can withdraw its entire body. Tubeworms are found among the following taxa: * Annelida, the ...
s, such as the
giant tube worm ''Riftia pachyptila'', commonly known as the giant tube worm and less commonly known as the Giant beardworm, is a marine invertebrate in the phylum Annelida (formerly grouped in phylum Pogonophora and Vestimentifera) related to tube worm ...
which lives in waters near underwater
volcanoes A volcano is a rupture in the crust of a planetary-mass object, such as Earth, that allows hot lava, volcanic ash, and gases to escape from a magma chamber below the surface. On Earth, volcanoes are most often found where tectonic plates a ...
and can withstand temperatures up to 90 degrees
Celsius The degree Celsius is the unit of temperature on the Celsius scale (originally known as the centigrade scale outside Sweden), one of two temperature scales used in the International System of Units (SI), the other being the Kelvin scale. The d ...
.
Platyhelminthes The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegm ...
(flatworms) form another worm phylum which includes a class
Cestoda Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of ...
of parasitic tapeworms. The marine tapeworm ''
Polygonoporus giganticus ''Tetragonoporus'' is a genus of cestodes in the order Pseudophyllidea. It is a monotypic genus, and the only species is ''Tetragonoporus calyptocephalus'', previously known as ''Polygonoporus giganticus''. This tapeworm is a gut parasite of wha ...
'', found in the gut of
sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...
s, can grow to over 30 m (100 ft).
Nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s (roundworms) constitute a further worm phylum with tubular
digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
s and an opening at both ends. Over 25,000 nematode species have been described, of which more than half are parasitic. It has been estimated another million remain undescribed. They are ubiquitous in marine, freshwater and terrestrial environments, where they often outnumber other animals in both individual and species counts. They are found in every part of the earth's
lithosphere A lithosphere () is the rigid, outermost rocky shell of a terrestrial planet or natural satellite. On Earth, it is composed of the crust and the portion of the upper mantle that behaves elastically on time scales of up to thousands of years ...
, from the top of mountains to the bottom of
oceanic trench Oceanic trenches are prominent long, narrow topographic depressions of the ocean floor. They are typically wide and below the level of the surrounding oceanic floor, but can be thousands of kilometers in length. There are about of oceanic tren ...
es. By count they represent 90% of all animals on the
ocean floor The seabed (also known as the seafloor, sea floor, ocean floor, and ocean bottom) is the bottom of the ocean. All floors of the ocean are known as 'seabeds'. The structure of the seabed of the global ocean is governed by plate tectonics. Most of ...
. * Their numerical dominance, often exceeding a million individuals per square meter and accounting for about 80% of all individual animals on earth, their diversity of life cycles, and their presence at various trophic levels point at an important role in many ecosystems. The bootlace worm can grow to . File:Riftia tube worm colony Galapagos 2011.jpg,
Giant tube worm ''Riftia pachyptila'', commonly known as the giant tube worm and less commonly known as the Giant beardworm, is a marine invertebrate in the phylum Annelida (formerly grouped in phylum Pogonophora and Vestimentifera) related to tube worm ...
s cluster around
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspo ...
s File:Lamellibrachia luymesi1.png, '' Lamellibrachia luymes'', a
cold seep A cold seep (sometimes called a cold vent) is an area of the ocean floor where hydrogen sulfide, methane and other hydrocarbon-rich fluid seepage occurs, often in the form of a brine pool. ''Cold'' does not mean that the temperature of the see ...
tubeworm, lives over 250 years. File:CSIRO ScienceImage 2818 Group of Nematodes.jpg,
Nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s are ubiquitous pseudocoelomates which can parasite marine plants and animals. File:Nerr0328.jpg, Bloodworms are typically found on the bottom of shallow marine waters File:Eunice aphroditois.jpg,
Bobbit worm ''Eunice aphroditois'' is a benthic bristle worm of warm marine waters. It lives mainly in the Atlantic Ocean, but can also be found in the Indo-Pacific. It ranges in length from less than to . Its iridescent cuticle produces a wide range of co ...
s are
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey ...
s that live on the seafloor


Echinoderms

Echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s (Greek for ''spiny skin'') is a phylum which contains only marine invertebrates. The adults are recognizable by their
radial symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
(usually five-point) and include
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish a ...
,
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) o ...
s,
sand dollar Sand dollars (also known as a sea cookie or snapper biscuit in New Zealand, or pansy shell in South Africa) are species of flat, burrowing sea urchins belonging to the order Clypeasteroida. Some species within the order, not quite as flat, are k ...
s, and
sea cucumber Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuri ...
s, as well as the
sea lilies Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
. Echinoderms are found at every ocean depth, from the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...
to the
abyssal zone The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word , meaning bottomless. At depths of , this zone remains in perpetual darkness. It covers 83% of the total area of the ocean an ...
. The phylum contains about 7000 living
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, making it the second-largest grouping of
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some ...
s (a superphylum), after the
chordates A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...
(which include the
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, such as
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s,
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
es,
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s, and
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 rhynchocephalia ...
s). Echinoderms are unique among animals in having bilateral symmetry at the larval stage, but fivefold symmetry ( pentamerism, a special type of radial symmetry) as adults. The echinoderms are important both biologically and geologically. Biologically, there are few other groupings so abundant in the biotic desert of the
deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of 200 metres (656 feet) or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combin ...
, as well as shallower oceans. Most echinoderms are able to regenerate tissue, organs, limbs, and reproduce asexually; in some cases, they can undergo complete regeneration from a single limb. Geologically, the value of echinoderms is in their
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s, which are major contributors to many
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms w ...
formations, and can provide valuable clues as to the geological environment. They were the most used species in regenerative research in the 19th and 20th centuries. Further, it is held by some scientists that the radiation of echinoderms was responsible for the Mesozoic Marine Revolution. File:Riccio Melone a Capo Caccia adventurediving.it.jpg,
Echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
literally means "spiny skin", as this water melon sea urchin illustrates File:Espardenya (animal).jpg,
Sea cucumber Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuri ...
s filter feed on plankton and suspended solids File:Enypniastes sp.jpg, Benthopelagic sea cucumbers can lift off the seafloor and journey as much as up the water column File:Ochre sea star.jpg, The
ochre sea star ''Pisaster ochraceus'', generally known as the purple sea star, ochre sea star, or ochre starfish, is a common seastar found among the waters of the Pacific Ocean. Identified as a keystone species, ''P. ochraceus'' is considered an important ind ...
was the first
keystone predator A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintain ...
to be studied. They limit
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which ...
s which can overwhelm intertidal communities.Holsinger, K. (2005). Keystone species. Retrieved May 10, 2010, from File:Colorful crinoids at shallow waters of Gili Lawa Laut.JPG, Colorful
sea lilies Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
in shallow waters
Aside from the hard-to-classify '' Arkarua'' (a
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of th ...
animal with echinoderm-like pentamerous radial symmetry), the first definitive members of the phylum appeared near the start of the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
.


Marine molluscs

Mollusc 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 est ...
s (Latin for ''soft'') form a
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
with about 85,000
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, ...
recognized
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
.Chapman, A.D. (2009)
Numbers of Living Species in Australia and the World, 2nd edition
Australian Biological Resources Study, Canberra. Retrieved 12 January 2010. (printed); (online).
By species count they are the largest
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military ...
phylum, comprising about 23% of all the named marine
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s. Molluscs have more varied forms than other invertebrate phyla. They are highly diverse, not just in size and in
anatomical Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
structure, but also in behaviour and in habitat. The majority of species still live in the oceans, from the seashores to the
abyssal zone The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word , meaning bottomless. At depths of , this zone remains in perpetual darkness. It covers 83% of the total area of the ocean an ...
, but some form a significant part of the freshwater fauna and the terrestrial
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s. The mollusc phylum is divided into 9 or 10 taxonomic classes, two of which are
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. These classes include
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s,
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
s and
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, as well as other lesser-known but distinctive classes.
Gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s with protective shells are referred to as
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
s (
sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the a ...
s), whereas gastropods without protective shells are referred to as
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a ...
s (
sea slug Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails (marine gastropod mollusks) that over evolutionary tim ...
s).
Gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s are by far the most numerous molluscs in terms of classified species, accounting for 80% of the total.
Bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
s include
clam Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shel ...
s,
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
s, cockles,
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which ...
s,
scallop Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
s, and numerous other families. There are about 8,000 marine bivalves species (including
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
and
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
species), and about 1,200 freshwater species.
Cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
include
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, ...
,
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fittin ...
and
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
. They are found in all oceans, and neurologically are the most advanced of the invertebrates. About 800 living species of marine cephalopods have been identified, and an estimated 11,000 extinct
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
have been described. There are no fully freshwater cephalopods. File:Calmarcolossal.jpg,
Colossal squid The colossal squid (''Mesonychoteuthis hamiltoni'') is part of the family Cranchiidae. It is sometimes called the Antarctic squid or giant cranch squid and is believed to be the largest squid species in terms of mass. It is the only recognize ...
, largest of all invertebrates File:Nautilus Palau.JPG, The
nautilus The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in ...
is a
living fossil A living fossil is an extant taxon that cosmetically 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 origin of the extant clade. Living foss ...
little changed since it evolved 500 million years ago as one of the first
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s. File:Nembrotha aurea B.jpg, Marine
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s are
sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the a ...
s or
sea slug Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails (marine gastropod mollusks) that over evolutionary tim ...
s. This
nudibranch Nudibranchs () are a group of soft-bodied marine gastropod molluscs which shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to match, ...
is a sea slug. File:Syrinx aruanus shell.jpg, The sea snail ''
Syrinx aruanus ''Syrinx aruanus'', common name the Australian trumpet or false trumpet, is a species of extremely large sea snail measuring up to 91 cm long and weighing up to 18 kg. It is a marine gastropod mollusk in the family Turbinellidae, ...
'' has the largest shell of any living gastropod File:Placopecten magellanicus.jpg, Molluscs usually have eyes. Bordering the edge of the mantle of a
scallop Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
, a
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
mollusc, can be over 100
simple eyes A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. File:Mytilus edulis illustration.png, Common mussel, another bivalve
Molluscs have such diverse shapes that many textbooks base their descriptions of molluscan anatomy on a generalized or ''hypothetical ancestral mollusc''. This generalized mollusc is unsegmented and
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...
with an underside consisting of a single muscular
foot The foot ( : feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg mad ...
. Beyond that it has three further key features. Firstly, it has a muscular cloak called a
mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
covering its viscera and containing a significant cavity used for breathing and
excretion Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after ...
. A
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 ** Thin-shell structure Science Biology * Seashell, a hard o ...
secreted by the mantle covers the upper surface. Secondly (apart from bivalves) it has a rasping tongue called a
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
used for feeding. Thirdly, it has a
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
including a complex digestive system using microscopic, muscle-powered hairs called
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proje ...
to exude
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
. The generalized mollusc has two paired nerve cords (three in bivalves). The
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
, in species that have one, encircles the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to t ...
. Most molluscs have
eyes Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and con ...
and all have sensors detecting chemicals, vibrations, and touch. The simplest type of molluscan
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are ...
relies on
external fertilization External fertilization is a mode of reproduction in which a male organism's sperm fertilizes a female organism's egg outside of the female's body. It is contrasted with internal fertilization, in which sperm are introduced via insemination and then ...
, but more complex variations occur. All produce
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 ...
, from which may emerge
trochophore A trochophore (; also spelled trocophore) is a type of free-swimming planktonic marine larva with several bands of cilia. By moving their cilia rapidly, they make a water eddy, to control their movement, and to bring their food closer, to captur ...
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. T ...
, more complex
veliger A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells. Description The veliger is the characteristic larva of the gastropod, bivalve and scaphopod ...
larvae, or miniature adults. The depiction is rather similar to modern
monoplacophora Monoplacophora , meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell inhabiting deep sea environments . Extant representatives were not recognized as such until 1952; previously they were known only from ...
ns, and some suggest it may resemble very early molluscs. Good evidence exists for the appearance of marine gastropods,
cephalopods A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, a ...
and bivalves in the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
period . However, the evolutionary history both of molluscs' emergence from the ancestral
Lophotrochozoa Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, bra ...
and of their diversification into the well-known living and
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
forms are still subjects of vigorous debate among scientists.


Marine arthropods

Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s (Greek for ''jointed feet'') have an
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
(external
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
), a segmented body, and jointed
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ante ...
s (paired appendages). They form a
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
which includes
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 pa ...
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 vinegar ...
s,
myriapod Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, ...
s, and
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s. Arthropods are characterized by their jointed limbs and
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
made of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, often mineralised with calcium carbonate. The arthropod
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
consists of segments, each with a pair of
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ante ...
s. The rigid cuticle inhibits growth, so arthropods replace it periodically by
moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
. Their versatility has enabled them to become the most species-rich members of all ecological guilds in most environments. Marine arthropods range in size from the microscopic
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
''
Stygotantulus ''Stygotantulus'' is a genus of crustacean with the sole species ''Stygotantulus stocki''. It lives as an ectoparasite on harpacticoid copepods of the families Tisbidae and Canuellidae. It is the smallest arthropod Arthropods (, (gen. ποδ ...
'' to the
Japanese spider crab The Japanese spider crab (''Macrocheira kaempferi'') is a species of marine crab that lives in the waters around Japan. It has the largest known leg-span of any arthropod. It goes through three main larval stages along with a prezoeal stage to gr ...
. Arthropods' primary internal cavity is a
hemocoel The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, t ...
, which accommodates their internal organs, and through which their
haemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
- analogue of
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the cir ...
- circulates; they have open circulatory systems. Like their exteriors, the internal organs of arthropods are generally built of repeated segments. Their
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
is "ladder-like", with paired
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
nerve cords running through all segments and forming paired
ganglia A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympathe ...
in each segment. Their heads are formed by fusion of varying numbers of segments, and their
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
s are formed by fusion of the ganglia of these segments and encircle the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to t ...
. The
respiratory 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 gre ...
and
excretory Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks aft ...
systems of arthropods vary, depending as much on their environment as on the
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of " subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists have also used th ...
to which they belong. Their vision relies on various combinations of
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s and pigment-pit
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
: in most species the ocelli can only detect the direction from which light is coming, and the compound eyes are the main source of information, but the main eyes of
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s are ocelli that can form images and, in a few cases, can swivel to track prey. Arthropods also have a wide range of chemical and mechanical sensors, mostly based on modifications of the many setae (bristles) that project through their cuticles. Arthropods' methods of reproduction and development are diverse; all terrestrial species use internal fertilization, but this is often by indirect transfer of the sperm via an appendage or the ground, rather than by direct injection. Marine species all lay eggs and use either internal or
external fertilization External fertilization is a mode of reproduction in which a male organism's sperm fertilizes a female organism's egg outside of the female's body. It is contrasted with internal fertilization, in which sperm are introduced via insemination and then ...
. Arthropod hatchlings vary from miniature adults to grubs that lack jointed limbs and eventually undergo a total metamorphosis to produce the adult form. File:Olenoides typicalis.jpg, Trilobites, now extinct, roamed oceans for 270 million years. File:Limule(dD) rotated.jpg, Horseshoe crab, a
living fossil A living fossil is an extant taxon that cosmetically 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 origin of the extant clade. Living foss ...
arthropod from 450 million years ago
File:Hyperia.jpg, Many crustaceans are very small, like this tiny amphipod, and make up a significant part of the ocean's zooplankton File:Stenopus hispidus (high res).jpg, The Stenopus hispidus, banded cleaner shrimp is a crustacean common in the tropics. File:J J Wild Pseudocarcinus cropped.jpg, The Tasmanian giant crab is long-lived and slow-growing, making it vulnerable to overfishing. File:Macrocheira kaempferi.jpg, The
Japanese spider crab The Japanese spider crab (''Macrocheira kaempferi'') is a species of marine crab that lives in the waters around Japan. It has the largest known leg-span of any arthropod. It goes through three main larval stages along with a prezoeal stage to gr ...
has the longest leg span of any arthropod.
The evolutionary ancestry of arthropods dates back to the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
period. The group is generally regarded as monophyletic, and many analyses support the placement of arthropods with cycloneuralians (or their constituent clades) in a superphylum Ecdysozoa. Overall however, the basal (evolution), basal relationships of Metazoa are not yet well resolved. Likewise, the relationships between various arthropod groups are still actively debated.


Other phyla

* Tardigrade, Lobopodia, (Onychophora) * Non-craniate (non-vertebrate) chordates: Cephalochordate, Tunicata and ''Haikouella''. These invertebrates are close relatives of the vertebrates. File:Amphioxus.png, The lancelet, a small translucent fish-like Cephalochordate, is the closest living invertebrate relative of the vertebrates.Lancelet (amphioxus) genome and the origin of vertebrates
''Ars Technica'', 19 June 2008.
File:Ascidian (Rhopalaea Crassa) (4 cm).png, Fluorescent-colored sea squirts, ''Rhopalaea crassa''. Tunicates may provide clues to vertebrate (and therefore human) ancestry. File:23 salpchain frierson odfw (8253212250).jpg, Salp chain File:Gill slits.png, Pharyngeal slits, Gill slits in an acorn worm (left) and tunicate (right)


Minerals from sea water

There are a number of marine invertebrates that use minerals that are present in the sea in such minute quantities that they were undetectable until the advent of spectroscopy. Vanadium is concentrated by some tunicates for use in their blood cells to a level ten million times that of the surrounding seawater. Other tunicates similarly concentrate niobium and tantalum. Lobsters use copper in their respiratory pigment hemocyanin, despite the proportion of this metal in seawater being minute. Although these elements are present in vast quantities in the ocean, their extraction by man is not economic.


Glass models

File:Anthomedusae - Lymnorea trieda.JPG File:Museum Histoire Naturelle Geneva Blaschka Aurelia aurita 21102014.jpg, ''Aurelia aurita'' File:Museum Histoire Naturelle Geneva Blaschka Cnidaria 21102014.jpg, Cnidarian File:Blaschka Natural History Museum Londres 28072013 2.jpg, Octopus File:Museum Histoire Naturelle Geneva Blaschka Glaucus atlanticus 21102014.jpg, ''Glaucus atlanticus'' File:BlaschkaSW (4).JPG, Sea anemone File:BlaschkaPisaCU (7).JPG


See also

* Marine life * Marine biology * Marine vertebrate * List of marine aquarium invertebrate species * Wiwaxia


Notes


References


Other references


List of Animal Phyla
{{aquatic ecosystem topics, expanded=marine Marine animals, Invertebrates