HOME

TheInfoList



OR:

Nemertea is 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 f ...
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 in ...
s 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 have patterns of yellow, orange, red and green coloration. The foregut, stomach and intestine run a little below the midline of the body, the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, de ...
is at the tip of the tail, and the mouth is under the front. A little above the gut is the rhynchocoel, a cavity which mostly runs above the midline and ends a little short of the rear of the body. All species have a
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
which lies in the rhynchocoel when inactive but everts to emerge just above the mouth to capture the animal's prey with venom. A highly extensible muscle in the back of the rhynchocoel pulls the proboscis in when an attack ends. A few species with stubby bodies
filter feed Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
and have suckers at the front and back ends, with which they attach to a
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ...
. The brain is a ring of four
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 ...
, positioned around the rhynchocoel near the animal's front end. At least a pair of
ventral nerve cord The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, in ...
s connect to the brain and run along the length of the body. Most nemerteans have various
chemoreceptors A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
, and on their heads some species have a number of pigment-cup
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-le ...
, which can detect light but can not form an image. Nemerteans respire through the skin. They have at least two
lateral Lateral is a geometric term of location which may refer to: Healthcare *Lateral (anatomy), an anatomical direction *Lateral cricoarytenoid muscle *Lateral release (surgery), a surgical procedure on the side of a kneecap Phonetics *Lateral cons ...
vessels which are joined at the ends to form a loop, and these and the rhynchocoel are filled with fluid. There is no heart, and the flow of fluid depends on contraction of muscles in the vessels and the body wall. To filter out soluble waste products,
flame cell A flame cell is a specialized excretory cell found in the simplest freshwater invertebrates, including flatworms, rotifers and nemerteans; these are the simplest animals to have a dedicated excretory system. Flame cells function like a kidney, re ...
s are embedded in the front part of the two lateral fluid vessels, and remove the wastes through a network of pipes to the outside. All nemerteans move slowly, using their external cilia to glide on surfaces on a trail of
slime Slime may refer to: Biology * Slime mold, a broad term often referring to roughly six groups of Eukaryotes * Biofilm, an aggregate of microorganisms in which cells adhere to each other and/or to a surface * Slimy (fish), also known as the pon ...
, while larger species use muscular waves to crawl, and some swim by dorso-ventral undulations. A few live in the open ocean while the rest find or make hiding places on the bottom. About a dozen species inhabit freshwater, mainly in the tropics and subtropics, and another dozen species live on land in cool, damp places. Most nemerteans are
carnivores 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 so ...
, feeding on
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
s,
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 she ...
s and
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s. Some species of nemerteans are
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
s, and a few live
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
ly inside the mantle cavity of
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 estim ...
s. In most species the sexes are separate, but all the freshwater species are
hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have sepa ...
. Nemerteans often have numerous temporary
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sperm ...
s (
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
or
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testost ...
), and build temporary gonoducts (ducts from which the ova or sperm are emitted) opening to a
gonopore A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female it is the ...
, one per gonad, when the ova and sperm are ready. The eggs are generally fertilised externally. Some species shed them into the water, and others protect their eggs in various ways. The fertilized egg divides by
spiral cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
and grows by determinate development, in which the fate of a cell can usually be predicted from its predecessors in the process of division. The embryos of most
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 ...
develop either directly to form juveniles (like the adult but smaller) or larvae that resemble the planulas of
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 th ...
. However, some form a pilidium larva, in which the developing juvenile has a gut which lies across the larva's body, and usually eats the remains of the larva when it emerges. The bodies of some species fragment readily, and even parts cut off near the tail can grow full bodies (antle ( fragmentation). Traditional
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. A ...
divides the phylum in two classes,
Anopla Anopla (for changes in taxonomy, see reference from 2019) has long been used as name for a class of marine worms of the phylum Nemertea, characterized by the absence of stylets on the proboscis, the mouth being below or behind the brain, and by ...
("unarmed" – their proboscises do not have a little dagger) with two orders, and
Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution and systematics The record of nemerteans is extremely sparse, as would be expected ...
("armed" with a dagger) also with two orders. However, it is now accepted that Anopla are
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 co ...
, as one order of Anopla is more closely related to Enopla than to the other order of Anopla. The phylum Nemertea itself is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, its main
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
being the rhynchocoel and eversible proboscis. Traditional taxonomy says that nemerteans are closely related to
flatworms 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, unsegment ...
, but both phyla are regarded as members of the
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, brachiop ...
, a very large clade, sometimes viewed as a superphylum that also includes
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 estim ...
s,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
s,
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...
s,
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 l ...
and many other
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's memb ...
s.


History

In 1555
Olaus Magnus Olaus Magnus (October 1490 – 1 August 1557) was a Swedish writer, cartographer, and Catholic ecclesiastic. Biography Olaus Magnus (a Latin translation of his birth name Olof Månsson) was born in Linköping in October 1490. Like his elder b ...
wrote of a marine worm which was apparently long ("40 cubits"), about the width of a child's arm, and whose touch made a hand swell.
William Borlase William Borlase (2 February 169631 August 1772), Cornish antiquary, geologist and naturalist. From 1722, he was Rector of Ludgvan, Cornwall, where he died. He is remembered for his works ''The Antiquities of Cornwall'' (1754; 2nd ed., 1769) ...
wrote in 1758 of a "sea long worm", and in 1770 Gunnerus wrote a formal description of this animal, which he called ''Ascaris longissima''. Its current name, ''
Lineus longissimus 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 ...
'', was first used in 1806 by Sowerby. In 1995, a total of 1,149 species had been described and grouped into 250 genera. Nemertea are named after the Greek sea-
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are typ ...
Nemertes, one of the daughters of
Nereus In Greek mythology, Nereus ( ; ) was the eldest son of Pontus (the Sea) and Gaia (the Earth), with Pontus himself being a son of Gaia. Nereus and Doris became the parents of 50 daughters (the Nereids) and a son ( Nerites), with whom Nereus li ...
and Doris. Alternative names for the phylum have included Nemertini, Nemertinea, and Rhynchocoela. The
Nemertodermatida Nemertodermatida is a class of Acoelomorpha, comprising 18 species of millimetre-sized turbellariform, mostly interstitial worms. Taxonomy The order ''Nemertodermatida'' contains two families with 6 genera. The high level of cryptic diversity ...
are a separate phylum, whose closest relatives appear to be the
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 of ...
.


Description


Body structure and major cavities

The typical nemertean body is very thin in proportion to its length. The smallest are a few millimeters long, most are less than , and several exceed . The longest animal ever found, at long, may be a specimen of ''
Lineus longissimus 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 ...
'', Ruppert, Fox and Barnes refer to a ''
Lineus longissimus 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 ...
'' long, washed ashore after a storm off
St Andrews St Andrews ( la, S. Andrea(s); sco, Saunt Aundraes; gd, Cill Rìmhinn) is a town on the east coast of Fife in Scotland, southeast of Dundee and northeast of Edinburgh. St Andrews had a recorded population of 16,800 , making it Fife's four ...
in Scotland. Other estimates are about . Zoologists find it extremely difficult to measure this species. For comparison: * The longest recorded
blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can b ...
was . * The
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23  million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is ...
s ''
Argentinosaurus ''Argentinosaurus'' is a genus of giant sauropod dinosaur that lived during the Late Cretaceous period in what is now Argentina. Although it is only known from fragmentary remains, ''Argentinosaurus'' is one of the largest known land animals of ...
'' and ''
Patagotitan ''Patagotitan'' is a genus of titanosaurian sauropod dinosaur from the Cerro Barcino Formation in Chubut Province, Patagonia, Argentina. The genus contains a single species known from multiple individuals: ''Patagotitan mayorum'', first announc ...
'' are estimated at approximately and respectively. * A specimen of the Arctic giant jellyfish '' Cyanea capillata arctica'' was long. Although ''L. longissimus'' is usually only a few millimeters wide. The bodies of most nemerteans can stretch a lot, up to 10 times their resting length in some species, but reduce their length to 50% and increase their width to 300% when disturbed. A few have relatively short but wide bodies, for example ''Malacobdella grossa'' is up to long and wide, and some of these are much less stretchy. Smaller nemerteans are approximately cylindrical, but larger species are flattened dorso-ventrally. Many have visible patterns in various combinations of yellow, orange, red and green. The outermost layer of the body has no
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 ...
, but consists of a ciliated and
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream ( endocrine gland) or into cavities inside the body or its outer surface ( exocrine gland). Structure ...
ular
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 ...
containing
rhabdite Rhabdites (from Greek, ''rhabdos'', rod) are rodlike structures in the cells of the epidermis or underlying parenchyma in certain turbellarians, and in the epidermis of nemertea Nemertea is a phylum of animals also known as ribbon worms or ...
s, which form the
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 is ...
in which the cilia glide. Each ciliated cell has many cilia and
microvilli Microvilli (singular: microvillus) are microscopic cellular membrane protrusions that increase the surface area for diffusion and minimize any increase in volume, and are involved in a wide variety of functions, including absorption, secretion, ce ...
. The outermost layer rests on a thickened
basement membrane The basement membrane is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between epithelial tissues including mesothelium and ...
, the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
. Next to the dermis are at least three layers of muscles, some circular and some longitudinal. The combinations of muscle types vary between the different classes, but these are not associated with differences in movement. Nemerteans also have dorso-ventral muscles, which flatten the animals, especially in the larger species. Inside the concentric tubes of these layers is
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every ...
, a kind of connective tissue. In
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
species this tissue is gelatinous and buoyant. The mouth is ventral and a little behind the front of the body. The foregut, stomach and intestine run a little below the midline of the body and the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, de ...
is at the tip of the tail. Above the gut and separated from the gut by mesenchyme is the rhynchocoel, a cavity which mostly runs above the midline and ends a little short of the rear of the body. The rhynchocoel of class
Anopla Anopla (for changes in taxonomy, see reference from 2019) has long been used as name for a class of marine worms of the phylum Nemertea, characterized by the absence of stylets on the proboscis, the mouth being below or behind the brain, and by ...
has an orifice a little to the front of the mouth, but still under the front of the body. In the other class,
Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution and systematics The record of nemerteans is extremely sparse, as would be expected ...
, the mouth and the front of the rhynchocoel share an orifice. The rhynchocoel is a
coelom The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...
, as it is lined by
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 ...
.


Proboscis and feeding

The
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
is an infolding of the body wall, and sits in the rhynchocoel when inactive. When muscles in the wall of the rhynchocoel compress the fluid inside, the pressure makes the proboscis jump inside-out to along a canal called the rhynchodeum and through an orifice, the proboscis pore. The proboscis has a muscle which attaches to the back of the rhynchocoel, and which can stretch up to 30 times its inactive length and then retract the proboscis. The proboscis of the class
Anopla Anopla (for changes in taxonomy, see reference from 2019) has long been used as name for a class of marine worms of the phylum Nemertea, characterized by the absence of stylets on the proboscis, the mouth being below or behind the brain, and by ...
exits from an orifice which is separate from the mouth, coils around the prey and immobilizes it by sticky, toxic secretions. The Anopla can attack as soon as they move into the range of the proboscis. Some Anopla have branched proboscises which can be described as "a mass of sticky spaghetti". The animal then draws its prey into its mouth. In most of the class
Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution and systematics The record of nemerteans is extremely sparse, as would be expected ...
, the proboscis exits from a common orifice of the rhynchocoel and mouth. A typical member of this class has a stylet, a
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
barb, with which the animal stabs the prey many times to inject toxins and digestive secretions. The prey is then swallowed whole or, after partial digestion, its tissues are sucked into the mouth. The stylet is attached about one-third of distance from the end of the
evert Evert is a Dutch and Swedish short form of the Germanic masculine name "Everhard" (alternative Eberhard).Evert
at the
ed proboscis, which extends only enough to expose the stylet. On either side of the active stylet are sacs containing back-up stylets to replace the active one as the animal grows or an active one is lost. Instead of one stylet, the Polystilifera have a pad that bears many tiny stylets, and these animals have separate orifices for the proboscis and mouth, unlike other Enopla. The Enopla can only attack after contacting the prey. Some nemerteans, such as ''L. longissimus'', absorb organic food in solution through their skins, which may make the long, slim bodies an advantage.
Suspension feeding Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
is found only among the specialized symbiotic bdellonemerteans, which have a proboscis but no stylet, and use suckers to attach themselves to
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biva ...
s.


Respiration and circulatory system

Nemerteans lack specialized
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
, and respiration occurs over the surface of the body, which is long and sometimes flattened. Like other animals with thick body walls, they use fluid circulation rather than
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
to move substances through their bodies. The circulatory system consists of the rhynchocoel and peripheral vessels, while their blood is contained in the main body cavity. The fluid in the rhynchocoel moves substances to and from the proboscis, and functions as a fluid
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 ...
in everting the proboscis and in burrowing. The vessels circulate fluid round the whole body and the rhynchocoel provides its own local circulation. The circulatory vessels are a system of coeloms. In the simplest type of circulatory system, two lateral vessels are joined at the ends to form a loop. However, many species have additional long-wise and cross-wise vessels. There is no heart nor pumping vessels, and the flow of fluid depends on contraction of both the vessels and the body wall's muscles. In some species, circulation is intermittent, and fluid ebbs and flows in the long-wise vessels. The fluid in the vessels is usually colorless, but in some species it contains cells that are yellow, orange, green or red. The red type contain
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyt ...
and carry oxygen, but the function of the other pigments is unknown.


Excretion

Nemertea use organs called
protonephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
to excrete soluble waste products, especially
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at sevent ...
ous by-products of cellular
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cel ...
. In nemertean protonephridia,
flame cell A flame cell is a specialized excretory cell found in the simplest freshwater invertebrates, including flatworms, rotifers and nemerteans; these are the simplest animals to have a dedicated excretory system. Flame cells function like a kidney, re ...
s which filter out the wastes are embedded in the front part of the two lateral fluid vessels. The flame cells remove the wastes into two collecting ducts, one on either side, and each duct has one or more
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores. Nephridia are homologous to nephrons ...
s through which the wastes exit. Semiterrestrial and freshwater nemerteans have many more flame cells than marines, sometimes thousands. The reason may be that
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
is more difficult in non-marine environments.


Nervous-system and senses

The central nervous-system consists of a brain and paired
ventral nerve cord The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, in ...
s that connect to the brain and run along the length of the body. The brain is a
ring Ring may refer to: * Ring (jewellery), a round band, usually made of metal, worn as ornamental jewelry * To make a sound with a bell, and the sound made by a bell :(hence) to initiate a telephone connection Arts, entertainment and media Film and ...
of four
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 ...
, masses of nerve cells, positioned round the rhynchocoel near its front end – while the brains of most
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's memb ...
invertebrates encircle the foregut. Most nemertean species have just one pair of nerve cords, many species have additional paired cords, and some species also have a dorsal cord. In some species the cords lie within the skin, but in most they are deeper, inside the muscle layers. The central nervous-system is often red or pink because it contains
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyt ...
. This stores
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as well ...
for peak activity or when the animal experiences anoxia, for example while
burrowing An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
in oxygen-free
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand a ...
s. Some species have paired cerebral organs, sacs whose only openings are to the outside. Others species have unpaired evertible organs on the front of their heads. Some have slits along the side of the head or grooves obliquely across the head, and these may be associated with paired cerebral organs. All of these are thought to be
chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
s, and the cerebral organs may also aiding
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
. Small pits in the epidermis appear to be sensors. On their head, some species have a number of pigment-cup
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-le ...
, which can detect light but not form an image. Most nemerteans have two to six ocelli, although some have hundreds. A few tiny species that live between grains of sand have
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in ''Xenoturbella''. The statocyst cons ...
s, which sense balance. '' Paranemertes peregrina'', which feeds on polychaetes, can follow the prey's trails of mucus, and find its burrow by backtracking along its own trail of mucus.


Movement

Nemerteans generally move slowly, though they have occasionally been documented to successfully prey on spiders or insects. Most nemerteans use their external cilia to glide on surfaces on a trail of
slime Slime may refer to: Biology * Slime mold, a broad term often referring to roughly six groups of Eukaryotes * Biofilm, an aggregate of microorganisms in which cells adhere to each other and/or to a surface * Slimy (fish), also known as the pon ...
, some of which is produced by glands in the head. Larger species use muscular waves to crawl, and some aquatic species swim by dorso-ventral undulations. Some species burrow by means of muscular
peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
, and have powerful muscles. Some species of the
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Monostilifera, whose proboscis have one active stylet, move by extending the proboscis, sticking it to an object and pulling the animal toward the object.


Reproduction and life-cycle

Larger species often break up when stimulated, and the fragments often grow into full individuals. Some species fragment routinely and even parts near the tail can grow full bodies. But this kind of extreme regeneration is restricted to only a few types of nemerteans, and is assumed to be a derived feature. All reproduce sexually, and most species are
gonochoric In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric. Gonochorism c ...
(the sexes are separate), but all the freshwater forms are
hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have sepa ...
. Nemerteans often have numerous temporary
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sperm ...
s (
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
or
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testost ...
), forming a row down each side of the body in the
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every ...
. Temporary gonoducts (ducts from which the
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
or
sperm 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, wh ...
are emitted), one per gonad, are built when the ova and sperm are ready. The eggs are generally fertilised externally. Some species shed them into the water, some lay them in a burrow or tube, and some protect them by
cocoon Cocoon may refer to: *Cocoon (silk), a pupal casing made by moth caterpillars and other insect larvae *Apache Cocoon, web development software * ''Cocoon'' (film), a 1985 science fiction-fantasy film **'' Cocoon: The Return'', 1988 sequel to ''Coco ...
s or
gelatinous Gelatin or gelatine (from la, gelatus meaning "stiff" or "frozen") is a translucent, colorless, flavorless food ingredient, commonly derived from collagen taken from animal body parts. It is brittle when dry and rubbery when moist. It may also ...
strings. Some
bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above, and the abyssopelagic below. The bathypelagi ...
(deep sea) species have
internal fertilization Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For intern ...
, and some of these are
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
, growing their
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm c ...
s in the female's body. The
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...
(fertilised egg) divides by
spiral cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
and grows by determinate development, in which the fate of a cell can usually be predicted from its predecessors in the process of division. The embryos of most
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 ...
develop either directly to form juveniles (like the adult but smaller) or to form planuliform
larva 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. Th ...
e. The planuliform larva stage may be short-lived and
lecithotrophic Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and m ...
("yolky") before becoming a juvenile, or may be planktotrophic, swimming for some time and eating prey larger than microscopic particles. However, many members of the order
Heteronemertea Heteronemertea is a monophyletic order of about 500 species of nemertean worm. It contains genera such as '' Lineus'' and '' Cerebratulus'', and includes the largest and most muscular nemerteans. Almost all heteronemerteans have three primary ...
and the
palaeonemertea Palaeonemertea is a class of primitive nemertean worm. It may be para- or polyphyletic, consisting of three to five clades and totalling about 100 species. These worms have several apparently simple features and, as their name suggests, they ...
n
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
Hubrechtiidae form a pilidium larva, which can capture
unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
and which Maslakova describes as like a
deerstalker A deerstalker is a type of cap that is typically worn in rural areas, often for hunting, especially deer stalking. Because of the cap's popular association with the fictional detective Sherlock Holmes, it has become stereotypical headgear for ...
cap with the ear flaps pulled down. It has a gut which lies across the body, a mouth between the "ear flaps", but no anus. A small number of
imaginal disc An imaginal disc is one of the parts of a holometabolous insect larva that will become a portion of the outside of the adult insect during the pupal transformation. Contained within the body of the larva, there are pairs of discs that will form ...
s form, encircling the
archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesenc ...
(developing gut) and coalesce to form the juvenile. When it is fully formed, the juvenile bursts out of the larva body and usually eats it during this catastrophic
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some insec ...
. This larval stage is unique in that there are no
Hox gene Hox genes, a subset of homeobox genes, are a group of related genes that specify regions of the body plan of an embryo along the head-tail axis of animals. Hox proteins encode and specify the characteristics of 'position', ensuring that the cor ...
s involved during development, which are only found in the juveniles developing inside the larvae. The species '' Paranemertes peregrina'' has been reported as having a life span of around 18 months.


Ecological significance

Most nemerteans are marine animals that burrow in sediments, lurk in crevices between shells, stones or the holdfasts of
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
or
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
animals. Some live deep in the open oceans, and have gelatinous bodies. Others build semi-permanent burrows lined with
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 is ...
or produce
cellophane Cellophane is a thin, transparent sheet made of regenerated cellulose. Its low permeability to air, oils, greases, bacteria, and liquid water makes it useful for food packaging. Cellophane is highly permeable to water vapour, but may be coat ...
-like tubes. Mainly in the tropics and subtropics, about 12 species appear in freshwater, and about a dozen species live on land in cool, damp places, for example under rotting logs. The terrestrial ''Argonemertes dendyi'' is a native of Australia but has been found in the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
, in Sao Miguel in the
Azores ) , motto =( en, "Rather die free than subjected in peace") , anthem= ( en, "Anthem of the Azores") , image_map=Locator_map_of_Azores_in_EU.svg , map_alt=Location of the Azores within the European Union , map_caption=Location of the Azores wi ...
, in
Gran Canaria Gran Canaria (, ; ), also Grand Canary Island, is the third-largest and second-most-populous island of the Canary Islands, an archipelago off the Atlantic coast of Northwest Africa which is part of Spain. the island had a population of that co ...
, and in a
lava tube A lava tube, or pyroduct, is a natural conduit formed by flowing lava from a volcanic vent that moves beneath the hardened surface of a lava flow. If lava in the tube empties, it will leave a cave. Formation A lava tube is a type of lava ...
at Kaumana on the
Island of Hawaii Hawaii ( ; haw, Hawaii ) is the largest island in the United States, located in the state of Hawaii. It is the southeasternmost of the Hawaiian Islands, a chain of volcanic islands in the North Pacific Ocean. With an area of , it has 63% of ...
. It can build a cocoon, which allows it to avoid desiccation while being transported, and it may be able to build populations quickly in new areas as it is a
protandrous Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular ...
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have s ...
. Another terrestrial
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencla ...
, '' Geonemertes'', is mostly found in
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologi ...
but has species in the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
, widely across the
Indo-Pacific The Indo-Pacific is a vast biogeographic region of Earth. In a narrow sense, sometimes known as the Indo-West Pacific or Indo-Pacific Asia, it comprises the tropical waters of the Indian Ocean, the western and central Pacific Ocean, and the ...
, in
Tristan da Cunha Tristan da Cunha (), colloquially Tristan, is a remote group of volcanic islands in the South Atlantic Ocean. It is the most remote inhabited archipelago in the world, lying approximately from Cape Town in South Africa, from Saint Helena ...
in the South Atlantic, in
Frankfurt Frankfurt, officially Frankfurt am Main (; Hessian: , " Frank ford on the Main"), is the most populous city in the German state of Hesse. Its 791,000 inhabitants as of 2022 make it the fifth-most populous city in Germany. Located on its ...
, in the Canary Islands, in
Madeira ) , anthem = ( en, "Anthem of the Autonomous Region of Madeira") , song_type = Regional anthem , image_map=EU-Portugal_with_Madeira_circled.svg , map_alt=Location of Madeira , map_caption=Location of Madeira , subdivision_type=Sovereign st ...
and in the Azores. '' Geonemertes pelaensis'' has been implicated in the decline of native arthropod species on the
Ogasawara Islands The Bonin Islands, also known as the , are an archipelago of over 30 subtropical and tropical islands, some directly south of Tokyo, Japan and northwest of Guam. The name "Bonin Islands" comes from the Japanese word ''bunin'' (an archaic read ...
, where it was introduced in the 1980s. Most are
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 so ...
s, feeding on
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
s,
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 she ...
s and
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s, and may kill annelids of about their own size. They sometimes take fish, both living and dead. Insects and
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, a ...
s are the only known prey of the two terrestrial species of ''Argonemertes''. A few nemerteans are
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
s, and these generally have good distance
chemoreception A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
("smell") and are not selective about their prey. A few species live
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
ly inside the mantle cavity of molluscs and feed on micro-organisms filtered out by the host. Near
San Francisco San Francisco (; Spanish for " Saint Francis"), officially the City and County of San Francisco, is the commercial, financial, and cultural center of Northern California. The city proper is the fourth most populous in California and 17th ...
the nemertean ''
Carcinonemertes errans ''Carcinonemertes errans'' is a ribbon worm in the family Carcinonemertidae. It lives in symbiosis with the Dungeness crab (''Metacarcinus magister''), consuming the crab's developing eggs. In 1980 it was implicated in the collapse of the Dunge ...
'' has consumed about 55% of the total egg production of its host, the
Dungeness crab The Dungeness crab (''Metacarcinus magister'') is a species of crab inhabiting eelgrass beds and water bottoms along the west coast of North America. It typically grows to across the carapace and is a popular seafood. Its common name comes fro ...
''Metacarcinus magister''. ''C. errans'' is considered a significant factor in the collapse of the dungeness crab fishery. Other coastal nemerteans have devastated
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 she ...
beds. The few predators on nemerteans include bottom-feeding fish, some sea birds, a few invertebrates including
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 ara ...
s, and other nemerteans. Nemerteans' skins secrete toxins that deter many predators, but some crabs may clean nemerteans with one claw before eating them. The American '' Cerebratulus lacteus'' and the South African ''Polybrachiorhynchus dayi'', both called "tapeworms" in their respective localities, are sold as fish bait.


Taxonomy

Traditional taxonomic classification has divided the group into two classes and four orders: * Class
Anopla Anopla (for changes in taxonomy, see reference from 2019) has long been used as name for a class of marine worms of the phylum Nemertea, characterized by the absence of stylets on the proboscis, the mouth being below or behind the brain, and by ...
("unarmed"). Includes animals with proboscis without stylet, and a mouth underneath and behind the brain. ** Order
Palaeonemertea Palaeonemertea is a class of primitive nemertean worm. It may be para- or polyphyletic, consisting of three to five clades and totalling about 100 species. These worms have several apparently simple features and, as their name suggests, they ...
. Comprises 100 marine species. Their body wall has outer circular and inner length-wise muscles. In addition, ''Carinoma tremaphoros'' has circular and inner length-wise muscles in the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
; the extra muscle layers seem to be needed for
burrowing An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
by
peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
. ** Order
Heteronemertea Heteronemertea is a monophyletic order of about 500 species of nemertean worm. It contains genera such as '' Lineus'' and '' Cerebratulus'', and includes the largest and most muscular nemerteans. Almost all heteronemerteans have three primary ...
. Comprises about 400 species. The majority are marine, but three are freshwater. Their body-wall muscles are disposed in four layers, alternately circular and length-wise starting from the outermost layer. The order includes the strongest swimmers. Two
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencl ...
have branched proboscises. * Class
Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution and systematics The record of nemerteans is extremely sparse, as would be expected ...
("armed"). All have stylets except order Bdellonemertea. Their mouth is located underneath and ahead of the brain. Their main nerve cords run inside body-wall muscles. ** Order Bdellonemertea. Includes seven species, of which six live as
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
s in the mantle of large
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 she ...
s and one in that of a freshwater snail. The hosts
filter feed Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
and all the hosts steal food from them. These nemerteans have short, wide bodies and have no stylets but have a sucking
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struct ...
and a posterior stucker, with which they move like inchworms. ** Order Hoplonemertea. Comprises 650 species. They live in
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
and
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
sea water, in freshwater and on land. They feed by commensalism and
parasitism 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 has c ...
, and are armed with stylet(s) *** Suborder Monostilifera. Includes 500 species with a single central stylet. Some use the stylet for locomotion as well as for capturing prey. *** Suborder Polystilifera. Includes about 100 pelagic and 50 benthic species. Their pads bear many tiny stylets. Recent molecular phylogenetic studies divided the group into two superclasses, three classes, and eight orders: *Superclass Pronemertea **Class
Palaeonemertea Palaeonemertea is a class of primitive nemertean worm. It may be para- or polyphyletic, consisting of three to five clades and totalling about 100 species. These worms have several apparently simple features and, as their name suggests, they ...
***Order Carinomiformes ***Order Tubulaniformes ***Order Archinemertea *Superclass Neonemertea **Class Pilidiophora ***Order Hubrechtiiformes ***Order
Heteronemertea Heteronemertea is a monophyletic order of about 500 species of nemertean worm. It contains genera such as '' Lineus'' and '' Cerebratulus'', and includes the largest and most muscular nemerteans. Almost all heteronemerteans have three primary ...
**Class Hoplonemertea (= Enopla) ***Order Polystilifera ***Order Monostilifera (includes Bdellonemertea) *''incertae sedis'' ** Order Arhynchonemertea (provisionally has been separated its own class Arhynchocoela in 1995)


Evolutionary history


Fossil record

As nemerteans are mostly soft-bodied, one would expect fossils of them to be extremely rare. Knaust (2010) reported nemertean fossils and traces from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
of Germany. One might expect the stylet of a nemertean to be fossilized, since it is made of the mineral
calcium phosphate The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are white ...
, but no fossilized stylets have been found. 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 ...
fossil '' Amiskwia'' 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 fos ...
has been classed as a nemertean, based on a resemblance to some unusual deep-sea swimming nemerteans, but few paleontologists accept this classification as the Burgess Shale fossils show no evidence of rhynchocoel nor intestinal caeca. Knaust & Desrochers (2019) reported fossils of
vermiform Vermiform (ˈvərməˌfôrm) describes something shaped like a worm. The expression is often employed in biology and anatomy to describe usually soft body parts or animals that are more or less tubular or cylindrical. The word root is Latin, ''ve ...
organisms with a wide range of morphologies occurring on bedding planes from the Late
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
(
Katian The Katian is the second stage of the Upper Ordovician. It is preceded by the Sandbian and succeeded by the Hirnantian Stage. The Katian began million years ago and lasted for about 7.8 million years until the beginning of the Hirnantian millio ...
) Vauréal Formation ( Canada). In the specimens preserving the anterior end of the body, this end is pointed or rounded, bearing a rhynchocoel with the proboscis, which is characteristic for nemerteans. The authors attributed these fossils to nemerteans and interpreted them as the oldest record of the group reported so far. However, Knaust & Desrochers cautioned that partly preserved putative nemertean fossils might ultimately turn out to be fossils of
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 more ...
ns or
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
s. It has been suggested that '' Archisymplectes'', one of the Pennsylvanian-age animals from Mazon Creek in northern and central
Illinois Illinois ( ) is a state in the Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolitan areas include, Peoria and Rockfo ...
, may be a nemertean. This fossil, however, only preserves the outline of the "worm", and there is no evidence of a proboscis, so there is no certainty that it represents a nemertean.


Within Nemertea

There is no doubt that the phylum Nemertea is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
(meaning that the phylum includes all and only descendants of one ancestor that was also a member of the phylum.RFB 2004 pp. 2-3 The
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
(trait shared by an ancestor and all its descendants, but not by other groups) include the eversible proboscis located in the rhynchocoel. While Ruppert, Fox and Barnes (2004) treat the Palaeonemertea as monophyletic, Thollesson and Norenburg (2003) regard them as
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 co ...
and basal (contains the ancestors of the more recent clades). The
Anopla Anopla (for changes in taxonomy, see reference from 2019) has long been used as name for a class of marine worms of the phylum Nemertea, characterized by the absence of stylets on the proboscis, the mouth being below or behind the brain, and by ...
("unarmed") represent an
evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of sp ...
of nemerteans without stylets (comprising the
Heteronemertea Heteronemertea is a monophyletic order of about 500 species of nemertean worm. It contains genera such as '' Lineus'' and '' Cerebratulus'', and includes the largest and most muscular nemerteans. Almost all heteronemerteans have three primary ...
and the Palaeonemerteans), while
Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution and systematics The record of nemerteans is extremely sparse, as would be expected ...
("armed") are monophyletic, but find that Palaeonemertea is doubly paraphyletic, having given rise to both the Heteronemertea and the Enopla. Ruppert, Fox and Barnes (2004) treat the Bdellonemertea as a clade separate from the Hoplonemertea, while Thollesson and Norenburg (2003) believe the Bdellonemertea are a part of the Monostilifera (with one active stylet), which are within the Hoplonemertea – which implies that "Enopla" and "Hoplonemertea" are synonyms for the same branch of the tree. The Polystilifera (with many tiny stylets) are monophyletic.


Relationships with other phyla

English-language writings have conventionally treated nemerteans as acoelomate bilaterians that are most closely related to flatworms (Platyhelminthes). These pre-
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
analyses emphasised as shared features: multiciliated (with multiple cilia per cell), glandular epidermis; rod-shaped secretory bodies or rhabdites; frontal glands or organs;
protonephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
; and acoelomate body organization. However, multiciliated epidermal cells and epidermal gland cells are also found in
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 ...
,
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 lo ...
,
Sipuncula 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 ...
, Annelida, Mollusca and other
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 ...
. The rhabdites of nemertea have a different structure from those of flatworms at the microscopic scale. The frontal glands or organs of flatworms vary a lot in structure, and similar structures appear in small marine annelids and entoproct larvae. The protonephridia of nemertea and flatworms are different in structure, and in position – the
flame cell A flame cell is a specialized excretory cell found in the simplest freshwater invertebrates, including flatworms, rotifers and nemerteans; these are the simplest animals to have a dedicated excretory system. Flame cells function like a kidney, re ...
s of nemertea are usually in the walls of the fluid vessels and are served by "drains" from which the wastes exit by a small number of tubes through the skin, while the flame cells of flatworms are scattered throughout the body. Rigorous comparisons show no synapomorphies of nemertean and platyhelminth nephridia. According to more recent analyses, in the development of nemertean embryos, ecto
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Embry ...
(outer part of the mesoderm, which is the layer in which most of the internal organs are built) is derived from cells labelled 3a and 3b, and endomesoderm (inner part of the mesoderm) is derived from the 4d cell. Some of the ectomesoderm in
annelids The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolog ...
,
echiuran 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 lo ...
s and
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 estim ...
s is derived from cells 3a and 3b, while the ectomesoderm of
polyclad The Polycladida represents a highly diverse clade of free-living marine flatworms. They are known from the littoral to the sublittoral zone (extending to the deep hot vents), and many species are common from coral reefs. Only a few species are ...
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, unsegment ...
s is derived from the 2b cell and
acoel 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 of ...
flatworms produce no ectomesoderm. In nemerteans the space between the epidermis and the gut is mainly filled by well-developed muscles embedded in noncellular connective tissue. This structure is similar to that found in larger flatworms such as
polyclad The Polycladida represents a highly diverse clade of free-living marine flatworms. They are known from the littoral to the sublittoral zone (extending to the deep hot vents), and many species are common from coral reefs. Only a few species are ...
s and
triclad A planarian is one of the many flatworms of the traditional class Turbellaria. It usually describes free-living flatworms of the order Tricladida (triclads), although this common name is also used for a wide number of free-living platyhelmi ...
s, but a similar structure of body-wall muscles embedded in noncellular connective tissue is widespread among the
Spiralia The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral cleavage ...
(animals in which the early cell divisions make a spiral pattern) such as
sipuncula 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 ...
ns,
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 lo ...
ns and many annelids. Nemerteans' affinities with Annelida (including Echiura, Pogonophora,
Vestimentifera Siboglinidae is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera (the giant tube worms). The family is composed of about 100 species of vermiform creatures which live in thin tubes buried ...
and perhaps Sipuncula) and Mollusca make the ribbon-worms members of
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, brachiop ...
, which include about half of the extant animal phyla. Lophotrochozoa groups: those animals that feed using a
lophophore The lophophore () is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata.Brachiopoda,
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 l ...
, Phoronida,
Entoprocta Entoprocta (), or Kamptozoa , is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that d ...
); phyla in which most members' embryos develop into
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 (for example Annelida and Mollusca); and some other phyla (such as Platyhelminthes, Sipuncula, Gastrotricha, Gnathostomulida, Micrognathozoa, Nemertea, Phoronida, Platyhelminthes and
Rotifera The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John Ha ...
). These groupings are based on
molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, which compares sections of organisms DNA and
RNA Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohyd ...
. While analyses by molecular phylogeny are confident that members of Lophotrochozoa are more closely related to each other than of non-members, the relationships between members are mostly unclear. Most
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's memb ...
phyla outside the Lophotrochozoa are members of
Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller phyla. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on phylogenetic tr ...
("animals that
molt 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 ...
"), which include
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 chitin, o ...
a,
Nematoda 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 broa ...
and
Priapulida Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertilit ...
. Most other
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 and ...
phyla are in the
Deuterostomia 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 exampl ...
, which include
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 sea l ...
and
Chordata 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 ...
. The
Acoelomorpha Acoelomorpha is a subphylum of very simple and small soft-bodied animals with planula-like features which live in marine or brackish waters. They usually live between grains of sediment, swimming as plankton, or crawling on other organisms, suc ...
, which are neither protostomes nor deuterostomes, are regarded as basal bilaterians.


See also

*'' Emplectonema neesii'' *''
Lineus longissimus 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 ...
'' *'' Parborlasia corrugatus''


Notes


References


External links


The Marine Biological Laboratory: Phylum Nemertea (Nemertinea, Nemertini, Rhynchocoela)Nemertea LifeDeskVideo of a Nemertea in Puget Sound
{{Authority control Animal phyla Extant Ordovician first appearances