Chaetognatha
Encyclopedia
Chaetognatha, meaning hair-jaws, and commonly known as arrow worms, are a phylum of predatory marine worms that are a major component of plankton
worldwide. About 20% of the known species are benthic, that is belonging to the lowest zone of the ocean, or benthic zone
, and can attach to algae and rocks. They are found in all marine waters, from surface tropical waters and shallow tide pools to the deep sea and polar regions. Most chaetognaths are transparent and are torpedo shaped, but some deep-sea species are orange. They range in size from 2 to 120 mm (0.078740157480315 to 4.7 in).
There are more than 120 modern species assigned to over 20 genera. Despite the limited diversity of species, the number of individuals is large.
. The body is divided into a distinct head, trunk, and tail. There are between four and fourteen hooked, grasping spines on each side of their head, flanking a hollow vestibule containing the mouth. The spines are used in hunting, and covered with a flexible hood arising from the neck region when the animal is swimming. All chaetognaths are carnivorous
, preying on other planktonic animals.
The trunk bears one or two pairs of lateral fins incorporating structures superficially similar to the fin rays of fish. Unlike those of vertebrates, however, these are composed of a thickened basement membrane
extending from the epidermis
, and they are not homologous
. An additional caudal fin covers the post-anal tail. Two chaetognath species, Caecosagitta macrocephala and Eukrohnia fowleri, have bioluminescent organs in their fins.,
Chaetognaths swim in short bursts using a dorso-ventral undulating body motion, where their tail fin assists with propulsion and the body fins for stabilization and steering. Some species are known to use the neurotoxin
tetrodotoxin
to subdue prey.
The body cavity lacks a peritoneum
, and therefore resembles the pseudocoel of animals such as nematode
s, but is divided into one compartment on each side of the trunk, and additional compartments inside the head and tail. Although they have a mouth with one or two rows of tiny teeth, compound eyes, and a nervous system, they have no respiratory or circulatory systems.
The mouth opens into a muscular pharynx
which contains glands to lubricate the passage of food. From here, a straight intestine runs the length of the trunk to an anus
just in front of the tail. The intestine is the primary site of digestion and includes a pair of diverticula
near the anterior end.
The nervous system is reasonably simple, consisting of a ganglion
ated nerve ring surrounding the pharynx. The dorsal ganglion is the largest, but nerves extend from all the ganglia along the length of the body. Chaetognaths have two compound eyes, each consisting of a number of pigment-cup ocelli fused together. In addition, there is a number of sensory bristles arranged in rows along the side of the body, where they probably perform a function similar to that of the lateral line
in fish. An additional, curved, band of sensory bristles lies over the head and neck.
Materials are moved about the body cavity by cilia. Waste materials are simply excreted through the skin and anus.
The arrow worm rhabdomeres are derived from microtubule
s, which in turn form conical bodies, which contain granules and thread structures. The cone body is derived from a cilium, and the microtubules of the rhabdomeres are 20 nm long and 50 nm wide.
and sperm
. Each animal possesses a pair of testes within the tail, and a pair of ovaries
in the posterior region of the main body cavity. Immature sperm are released from the testes to mature inside the cavity of the tail, and then swim through a short duct to a seminal vesicle
where they are packaged into a spermatophore
.
During mating, each individual places a spermatophore onto the neck of its partner after rupture of the seminal vesicle. The sperm rapidly escape from the spermatophore and swim along the midline of the animal until they reach a pair of small pores just in front of the tail. These pores connect to the oviduct
s, into which the developed eggs have already passed from the ovaries, and it is here that fertilisation takes place.
The eggs are planktonic, or attached to nearby algae, and hatch into miniature versions of the adult, without a well-defined larva
l stage.
and K. V. Schwartz place chaetognaths in the deuterostomes in their Five Kingdom classification. Molecular phylogenists, however, consider them to be protostomes. Thomas Cavalier-Smith
places them in the protostomes in his Six Kingdom classification. The similarities between chaetognaths and nematodes mentioned above may support the protostome thesis - in fact, chaetognaths are sometimes regarded as a basal ecdysozoa
n or lophotrochozoa
n.
Chaetognatha appears close to the base of the protostome tree in most studies of their molecular phylogeny. This may explain their deuterostome embryonic characters. If chaetognaths branched off from the protostomes before they evolved their distinctive protostome embryonic characters, they may have retained deuterostome characters inherited from early bilateria
n ancestors. Thus chaetognaths may be a useful model for the ancestral bilaterian.
Studies of arrow worms' nervous systems suggests they should be placed within the protostomes.
Period. Complete body fossils have been formally described from the Lower Cambrian
Maotianshan shales
of Yunnan, China (Eognathacantha ercainella Chen & Huang and Protosagitta spinosa Hu) and the Middle Cambrian Burgess Shale
of British Columbia (Oesia disjuncta Walcott). A more recent chaetognath, Paucijaculum samamithion Schram, has been described from the Mazon Creek biota from the Pennsylvanian
of Illinois. Chaetognaths were thought possibly to be related to some of the animals grouped with the conodont
s. The conodonts themselves, however, are thought to be related to the vertebrates. It is now thought that protoconodont elements (e.g., Protohertzina anabarica Missarzhevsky, 1973), are probably grasping spines of chaetognaths rather than teeth of conodonts. Previously chaetognaths in the Early Cambrian were only suspected from these protoconodont elements, but the more recent discoveries of body fossils have confirmed their presence then.
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
worldwide. About 20% of the known species are benthic, that is belonging to the lowest zone of the ocean, or benthic zone
Benthic zone
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean or a lake, including the sediment surface and some sub-surface layers. Organisms living in this zone are called benthos. They generally live in close relationship with the substrate bottom; many such...
, and can attach to algae and rocks. They are found in all marine waters, from surface tropical waters and shallow tide pools to the deep sea and polar regions. Most chaetognaths are transparent and are torpedo shaped, but some deep-sea species are orange. They range in size from 2 to 120 mm (0.078740157480315 to 4.7 in).
There are more than 120 modern species assigned to over 20 genera. Despite the limited diversity of species, the number of individuals is large.
Anatomy
Chaetognaths are transparent or translucent dart-shaped animals covered by a cuticleCuticle
A cuticle , or cuticula, is a term used for 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 "cuticles" are non-homologous; differing in their origin, structure, function, and chemical composition...
. The body is divided into a distinct head, trunk, and tail. There are between four and fourteen hooked, grasping spines on each side of their head, flanking a hollow vestibule containing the mouth. The spines are used in hunting, and covered with a flexible hood arising from the neck region when the animal is swimming. All chaetognaths are carnivorous
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...
, preying on other planktonic animals.
The trunk bears one or two pairs of lateral fins incorporating structures superficially similar to the fin rays of fish. Unlike those of vertebrates, however, these are composed of a thickened basement membrane
Basement membrane
The basement membrane is a thin sheet of fibers that underlies the epithelium, which lines the cavities and surfaces of organs including skin, or the endothelium, which lines the interior surface of blood vessels.- Composition :...
extending from the epidermis
Epidermis (zoology)
The Epidermis is an epithelium that covers the body of an eumetazoan . Eumetazoa have a cavity lined with a similar epithelium, the gastrodermis, which forms a boundary with the epidermis at the mouth.Sponges have no epithelium, and therefore no epidermis or gastrodermis...
, and they are not homologous
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
. An additional caudal fin covers the post-anal tail. Two chaetognath species, Caecosagitta macrocephala and Eukrohnia fowleri, have bioluminescent organs in their fins.,
Chaetognaths swim in short bursts using a dorso-ventral undulating body motion, where their tail fin assists with propulsion and the body fins for stabilization and steering. Some species are known to use the neurotoxin
Neurotoxin
A neurotoxin is a toxin that acts specifically on nerve cells , usually by interacting with membrane proteins such as ion channels. Some sources are more general, and define the effect of neurotoxins as occurring at nerve tissue...
tetrodotoxin
Tetrodotoxin
Tetrodotoxin, also known as "tetrodox" and frequently abbreviated as TTX, sometimes colloquially referred to as "zombie powder" by those who practice Vodou, is a potent neurotoxin with no known antidote. There have been successful tests of a possible antidote in mice, but further tests must be...
to subdue prey.
The body cavity lacks a peritoneum
Peritoneum
The peritoneum is the serous membrane that forms the lining of the abdominal cavity or the coelom — it covers most of the intra-abdominal organs — in amniotes and some invertebrates...
, and therefore resembles the pseudocoel of animals such as nematode
Nematode
The nematodes or roundworms are the most diverse phylum of pseudocoelomates, and one of the most diverse of all animals. Nematode species are very difficult to distinguish; over 28,000 have been described, of which over 16,000 are parasitic. It has been estimated that the total number of nematode...
s, but is divided into one compartment on each side of the trunk, and additional compartments inside the head and tail. Although they have a mouth with one or two rows of tiny teeth, compound eyes, and a nervous system, they have no respiratory or circulatory systems.
The mouth opens into a muscular pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
which contains glands to lubricate the passage of food. From here, a straight intestine runs the length of the trunk to an anus
Anus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
just in front of the tail. The intestine is the primary site of digestion and includes a pair of diverticula
Diverticulum
A diverticulum is medical or biological term for an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, they are described as being either true or false....
near the anterior end.
The nervous system is reasonably simple, consisting of a ganglion
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
ated nerve ring surrounding the pharynx. The dorsal ganglion is the largest, but nerves extend from all the ganglia along the length of the body. Chaetognaths have two compound eyes, each consisting of a number of pigment-cup ocelli fused together. In addition, there is a number of sensory bristles arranged in rows along the side of the body, where they probably perform a function similar to that of the lateral line
Lateral line
The lateral line is a sense organ in aquatic organisms , used to detect movement and vibration in the surrounding water. Lateral lines are usually visible as faint lines running lengthwise down each side, from the vicinity of the gill covers to the base of the tail...
in fish. An additional, curved, band of sensory bristles lies over the head and neck.
Materials are moved about the body cavity by cilia. Waste materials are simply excreted through the skin and anus.
The arrow worm rhabdomeres are derived from microtubule
Microtubule
Microtubules are a component of the cytoskeleton. These rope-like polymers of tubulin can grow as long as 25 micrometers and are highly dynamic. The outer diameter of microtubule is about 25 nm. Microtubules are important for maintaining cell structure, providing platforms for intracellular...
s, which in turn form conical bodies, which contain granules and thread structures. The cone body is derived from a cilium, and the microtubules of the rhabdomeres are 20 nm long and 50 nm wide.
Reproduction
All species are hermaphroditic, carrying both eggsOvum
An ovum is a haploid female reproductive cell or gamete. Both animals and embryophytes have ova. The term ovule is used for the young ovum of an animal, as well as the plant structure that carries the female gametophyte and egg cell and develops into a seed after fertilization...
and sperm
Spermatozoon
A spermatozoon is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote...
. Each animal possesses a pair of testes within the tail, and a pair of ovaries
Ovary
The ovary is an ovum-producing reproductive organ, often found in pairs as part of the vertebrate female reproductive system. Ovaries in anatomically female individuals are analogous to testes in anatomically male individuals, in that they are both gonads and endocrine glands.-Human anatomy:Ovaries...
in the posterior region of the main body cavity. Immature sperm are released from the testes to mature inside the cavity of the tail, and then swim through a short duct to a seminal vesicle
Seminal vesicle
The seminal vesicles or vesicular glands are a pair of simple tubular glands posteroinferior to the urinary bladder of male mammals...
where they are packaged into a spermatophore
Spermatophore
A spermatophore or sperm ampulla is a capsule or mass created by males of various animal species, containing spermatozoa and transferred in entirety to the female's ovipore during copulation...
.
During mating, each individual places a spermatophore onto the neck of its partner after rupture of the seminal vesicle. The sperm rapidly escape from the spermatophore and swim along the midline of the animal until they reach a pair of small pores just in front of the tail. These pores connect to the oviduct
Oviduct
In non-mammalian vertebrates, the passageway from the ovaries to the outside of the body is known as the oviduct. The eggs travel along the oviduct. These eggs will either be fertilized by sperm to become a zygote, or will degenerate in the body...
s, into which the developed eggs have already passed from the ovaries, and it is here that fertilisation takes place.
The eggs are planktonic, or attached to nearby algae, and hatch into miniature versions of the adult, without a well-defined larva
Larva
A larva 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...
l stage.
Classification
Chaetognaths are traditionally classed as deuterostomes by embryologists. Lynn MargulisLynn Margulis
Lynn Margulis was an American biologist and University Professor in the Department of Geosciences at the University of Massachusetts Amherst. She is best known for her theory on the origin of eukaryotic organelles, and her contributions to the endosymbiotic theory, which is now generally accepted...
and K. V. Schwartz place chaetognaths in the deuterostomes in their Five Kingdom classification. Molecular phylogenists, however, consider them to be protostomes. Thomas Cavalier-Smith
Thomas Cavalier-Smith
Professor Thomas Cavalier-Smith , FRS, FRSC, NERC Professorial Fellow, is a Professor of Evolutionary Biology in the Department of Zoology, at the University of Oxford...
places them in the protostomes in his Six Kingdom classification. The similarities between chaetognaths and nematodes mentioned above may support the protostome thesis - in fact, chaetognaths are sometimes regarded as a basal ecdysozoa
Ecdysozoa
Ecdysozoa is a group of protostome animals, including Arthropoda , Nematoda, and several smaller phyla. They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes...
n or lophotrochozoa
Lophotrochozoa
The Lophotrochozoa are a major grouping of protostome animals. The taxon was discovered based on molecular data. Molecular evidence such as a result of studies of the evolution of small-subunit ribosomal RNA supports the monophyly of the phyla listed in the infobox shown at right.-Terminology:The...
n.
Chaetognatha appears close to the base of the protostome tree in most studies of their molecular phylogeny. This may explain their deuterostome embryonic characters. If chaetognaths branched off from the protostomes before they evolved their distinctive protostome embryonic characters, they may have retained deuterostome characters inherited from early bilateria
Bilateria
The bilateria are all animals having a bilateral symmetry, i.e. they have a front and a back end, as well as an upside and downside. Radially symmetrical animals like jellyfish have a topside and downside, but no front and back...
n ancestors. Thus chaetognaths may be a useful model for the ancestral bilaterian.
Studies of arrow worms' nervous systems suggests they should be placed within the protostomes.
Fossil record
Due to their soft bodies, chaetognaths fossilize poorly. Even so, several fossil chaetognath species have been described. Chaetognaths appear to have originated in the CambrianCambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
Period. Complete body fossils have been formally described from the Lower Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
Maotianshan shales
Maotianshan shales
The Maotianshan Shales are a series of lower Cambrian deposits in the Chiungchussu formation, famous for their Konservat Lagerstätten, or high number of fossils preserved in place...
of Yunnan, China (Eognathacantha ercainella Chen & Huang and Protosagitta spinosa Hu) and the Middle Cambrian Burgess Shale
Burgess Shale
The Burgess Shale Formation, located in the Canadian Rockies of British Columbia, is one of the world's most celebrated fossil fields, and the best of its kind. It is famous for the exceptional preservation of the soft parts of its fossils...
of British Columbia (Oesia disjuncta Walcott). A more recent chaetognath, Paucijaculum samamithion Schram, has been described from the Mazon Creek biota from the Pennsylvanian
Pennsylvanian
The Pennsylvanian is, in the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly . As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain...
of Illinois. Chaetognaths were thought possibly to be related to some of the animals grouped with the conodont
Conodont
Conodonts are extinct chordates resembling eels, classified in the class Conodonta. For many years, they were known only from tooth-like microfossils now called conodont elements, found in isolation. Knowledge about soft tissues remains relatively sparse to this day...
s. The conodonts themselves, however, are thought to be related to the vertebrates. It is now thought that protoconodont elements (e.g., Protohertzina anabarica Missarzhevsky, 1973), are probably grasping spines of chaetognaths rather than teeth of conodonts. Previously chaetognaths in the Early Cambrian were only suspected from these protoconodont elements, but the more recent discoveries of body fossils have confirmed their presence then.
