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Jaw worms
Gnathostomula paradoxa Sylt.tif
Gnathostomula paradoxa
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Clade: Gnathifera
Phylum: Gnathostomulida
Ax, 1956 [1]
Orders and suborders

Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956. [1]



Most gnathostomulids measure 0.5 to 1 millimetre (0.02 to 0.04 in) in length. They are often slender to thread-like worms, with a generally transparent body. In many Bursovaginoidea, one of the major group of gnathostomulids, the neck region is slightly narrower than the rest of the body, giving them a distinct head. [2]

Like flatworms they have a ciliated epidermis, but in contrast to flatworms, they have one cilium per cell. [3] The cilia allow the worms to glide along in the water between sand grains, although they also use muscles, allowing the body to twist or contract, for movement.

They have no body cavity, and no circulatory or respiratory system. The nervous system is simple, and restricted to the outer layers of the body wall. The only sense organs are modified cilia, which are especially common in the head region. [2]

The mouth is located just behind the head, after a rostrum, on the underside of the body. It has a pair of cuticular jaws, supplied by strong muscles, and often bearing minute teeth. A "basal plate" on the lower surface that bears a comb-like structure is also present. The basal plate is used to scrape smaller organisms off of the grains of sand that make up their anoxic seabed mud habitat. [4] This bilaterally symmetrical pharynx with its complex cuticular mouth parts make them appear closely related to rotifers and their allies, together making up the Gnathifera. The ultrastructure of the jaws made of rods with electron dense core in transmission electron microscopy sections also support their close relation with Rotifera and Micrognathozoa. The mouth opens into a blind-ending tube in which digestion takes place; there is no true anus. [2] However, there is tissue connecting the intestine to the epidermis which may serve as an anal pore. [5]


Gnathostomulids are simultaneous hermaphrodites. Each individual possesses a single ovary and one or two testes. After fertilization, the single egg ruptures through the body wall and adheres to nearby sand particles; the parent is able to rapidly heal the resulting wound. The egg hatches into a miniature version of the adult, without a larval stage. [2]


There are approximately 100 described species [6] and certainly many more as yet undescribed. The known species are grouped in two orders. The filospermoids are very long and are characterized by an elongate rostrum. The bursovaginoids have paired sensory organs and are characterized by the presence of a penis and a sperm-storage organ called a bursa. [4]

Gnathostomulids have no known fossil record, though there are (debatable) similarities between the jaws of modern gnathostomulids and certain conodont elements. (Ochietti & Cailleux, 1969; Durden et al, 1969) [7]

They appear to be a sister clade to the Syndermata. [2] [8]

Related Research Articles

Sipuncula Phylum of invertebrates, peanut worms

The Sipuncula or Sipunculida is a group containing about 162 species of bilaterally symmetrical, unsegmented marine worms. The name Sipuncula is from the genus name Sipunculus, and comes from the Latin siphunculus meaning a "small tube". Traditionally considered a phylum, Sipuncula seems to be closely related to Myzostomida and Annelida, and may be a subgroup of Annelida, based on recent molecular work. Sipunculans vary in size but most species are under 10 cm (4 in) in length.

Gastrotrich Phylum of microscopic pseudocoelomate animals

The gastrotrichs, commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, pseudocoelomate animals, and are widely distributed and abundant in freshwater and marine environments. They are mostly benthic and live within the periphyton, the layer of tiny organisms and detritus that is found on the seabed and the beds of other water bodies. The majority live on and between particles of sediment or on other submerged surfaces, but a few species are terrestrial and live on land in the film of water surrounding grains of soil. Gastrotrichs are divided into two orders, the Macrodasyida which are marine, and the Chaetonotida, some of which are marine and some freshwater. Nearly 800 species of gastrotrich have been described.

Onychophora Phylum of invertebrates, velvet worms

Onychophora, commonly known as velvet worms or more ambiguously as peripatus, is a phylum of elongate, soft-bodied, many-legged panarthropods. In appearance they have variously been compared to worms with legs, caterpillars, and slugs. They prey upon smaller animals such as insects, which they catch by squirting an adhesive slime.

Nemertea Phylum of invertebrates, ribbon worms

Nemertea is a phylum of invertebrate animals also known as ribbon worms or proboscis worms. Alternative names for the phylum have included Nemertini, Nemertinea and Rhynchocoela. Most 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.

Bilateria All animals having a bilateral symmetry as embryo

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 as well as a back and a belly. Nearly all are bilaterally symmetrical as adults as well; the most notable exception is the echinoderms, which achieve secondary pentaradial symmetry as adults, but are bilaterally symmetrical during embryonic development.

Coelom The main body cavity in most animals

The coelom 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 remains undifferentiated.

Entoprocta A phylum of aquatic invertebrates

Entoprocta whose name means "anus inside", or Kamptozoa, is a phylum of mostly sessile aquatic animals, ranging from 0.1 to 7 millimetres long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and both the mouth and anus lie inside the "crown". The superficially similar Bryozoa (Ectoprocta) have the anus outside a "crown" of hollow tentacles. Most families of entoprocts are colonial, and all but 2 of the 150 species are marine. A few solitary species can move slowly.

Planarian one of many flatworms of the Turbellaria class. It is also the common name for a member of the genus Planaria within the family Planariidae. Sometimes it also refers to the genus Dugesia

A planarian is one of many flatworms of the 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 platyhelminthes. Planaria are common to many parts of the world, living in both saltwater and freshwater ponds and rivers. Some species are terrestrial and are found under logs, in or on the soil, and on plants in humid areas.

Turbellaria class of worms

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 1 mm (0.039 in) to large freshwater forms more than 500 mm (20 in) long or terrestrial species like Bipalium kewense which can reach 600 mm (24 in) in length. All the larger forms are flat with ribbon-like or leaf-like shapes, since their lack of respiratory and circulatory systems means that they have to rely on diffusion for internal transport of metabolites. However, many of the smaller forms are round in cross section. Most are predators, and all live in water or in moist terrestrial environments. Most forms reproduce sexually and with few exceptions all are simultaneous hermaphrodites.

<i>Limnognathia</i> monotypic taxon

Limnognathia maerski is a microscopic platyzoan freshwater animal, discovered living in warm springs on Disko Island, Greenland in 1994, that has variously been assigned as a class or subphylum in the phylum Gnathifera or as a phylum in a Gnathifera superphylum, named Micrognathozoa. It is related to the rotifers and gnathostomulids, grouped together as the Gnathifera. With an average length of one-tenth of a millimetre, it is one of the smallest animals known.

Platyzoa superphylum of animals

The paraphyletic "Platyzoa" are a group of protostome unsegmented animals proposed by Thomas Cavalier-Smith in 1998. Cavalier-Smith included in Platyzoa the phylum Platyhelminthes, and a new phylum, the Acanthognatha, into which he gathered several previously described phyla of microscopic animals. More recently it has been described as paraphyletic, containing the Rouphozoa and the Gnathifera.

Acorn worm Class of hemichordate invertebrates

The acorn worms or Enteropneusta are a hemichordate class of invertebrates consisting of one order of the same name. Their closest relatives are the echinoderms. There are 111 known species of acorn worm in the world, the main species for research being Saccoglossus kowalevskii. Two families - Harrimaniidae and Ptychoderidae - separated at least 370 million years ago.

Acoela order of worms

The Acoela or acoels are an order of small and simple invertebrates in the phylum Xenacoelomorpha, a basal Bilaterian group, which resemble flatworms. Historically they were treated as an order of turbellarian flatworms.

Marine invertebrates

Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.

Phoronid phylum of marine animals, horseshoe worms

Phoronids are a small phylum of marine animals that filter-feed with a lophophore, and build upright tubes of chitin to support and protect their soft bodies. They live in most of the oceans and seas including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about 400 meters down. Most adult phoronids are 2 cm long and about 1.5 mm wide, although the largest are 50 cm long.

Gnathifera is a clade of generally small spiralians characterized by complex jaws made of chitin. It comprises the phyla Gnathostomulida, Rotifera, Micrognathozoa, and Chaetognatha. It may also include the Cycliophora.

Listriolobus pelodes is a species of marine spoon worm. It is found in shallow seas in the North East Pacific off the coast of California. It lives in a burrow in soft sediments.

Annelid Phylum of segmented worms

The annelids, also known as the ringed worms or 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 ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.

<i>Tubulanus polymorphus</i> species of worm

Tubulanus polymorphus is a species of ribbon worm in the phylum Nemertea. It is found in the northern Atlantic Ocean and the northern Pacific Ocean. It occurs on the lower shore down to about 50 m (160 ft), on sand or gravel, under stones and among seaweed.

<i>Tubulanus superbus</i> species of worm

Tubulanus superbus, commonly known as the football jersey worm, is a species of ribbon worm in the phylum Nemertea. Found in the northern Atlantic Ocean, the North Sea and the Mediterranean Sea, it occurs from the lower shore down to about 80 m (260 ft), on sand or gravel.


  1. 1 2 Ax, P. (1956). "Die Gnathostomulida, eine rätselhafte Wurmgruppe aus dem Meeressand". Abhandl. Akad. Wiss. U. Lit. Mainz, Math. - Naturwiss. 8: 1–32.
  2. 1 2 3 4 5 Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 311–312. ISBN   0-03-056747-5.
  3. Ruppert, Edward E., Fox, Richard S., Barnes, Robert D. (2004) Invertebrate Zoology (7th edition). Brooks/Cole-Thomson Learning, Belmont, US
  4. 1 2 Barnes, R.F.K. et al. (2001). The Invertebrates: A Synthesis. Oxford: Blackwell Science.
  5. Knauss, Elizabeth (December 1979). "Indication of an Anal Pore in Gnathostomulida". Zoologica Scripta. 8 (1–4): 181–6. doi:10.1111/j.1463-6409.1979.tb00630.x.
  6. Zhang, Z.-Q. (2011). "Animal biodiversity: An introduction to higher-level classification and taxonomic richness" (PDF). Zootaxa. 3148: 7–12. doi:10.11646/zootaxa.3148.1.3.
  7. Cited in chapter 2 , p192 of Andreas Schmidt-Rhaesa'a 2015 Gastrotricha and Gnathifera, VOl 3 of Gast---, Cycloneuralia and Gnath---.
  8. Golombek, A.; Tobergte, S.; Struck, T.H. (May 2015). "Elucidating the phylogenetic position of Gnathostomulida and first mitochondrial genomes of Gnathostomulida, Gastrotricha and Polycladida (Platyhelminthes)". Mol Phylogenet Evol. 86: 49–63. doi:10.1016/j.ympev.2015.02.013. PMID   25796325.