Gnathostomulida (Gnathostomulids)

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Gnathostomulida

(Gnathostomulids)

Phylum Gnathostomulida

Number of families 12

Thumbnail description
Marine group of microscopic free-living worms characterized by an entirely monociliated epidermis and complex cuticular mouthparts


Evolution and systematics

There is no fossil record for this group. Described in 1956 as aberrant Turbellaria (flatworms), the gnathostomulids are now considered related to the rotifers (Rotifera) and Micrognathozoa. The phylum (and single class) encompasses two orders, Filospermoidea and Bursovaginoidea, the latter with two suborders, Conophoralia and Scleroperalia. The 12 families contain 25 genera, with fewer than 100 valid species.

Physical characteristics

These thread-shaped worms range from 0.01 in (0.3 mm) to more than 0.1 in (3 mm) in length. Most species are colorless or transparent, but a few are bright red. The anterior end of Filospermoidea is pointed, while that of Bursovaginoidea appears as a rounded head. The posterior end is rounded or extends into a tail. The epidermis is completely monociliated, i.e., each cell carries a single, long locomotory cilium. Some cilia, singly or in paired groups, may have sensory functions. The nervous system, which is largely situated at the basis of the epidermis, consists of an unpaired frontal ganglion (brain) and an unpaired buccal ganglion from which paired nerves originate. The musculature is simple and rather weak, except for a complex pharynx. Circulatory and respiratory organs are lacking. The digestive tract provides the greatest number of distinguishing characters. The mouth is located ventrally, behind the anterior end, and there is no permanent anus. In the majority of species the complex, muscular, bilaterally symmetric pharynx contains cuticular, hard mouthparts consisting of an unpaired basal plate in the lower lip, and paired jaws. The basal plate may be flat and dorsally set with ridges or teeth, as in the family Haplognathiidae; transverse rod shaped, as in Pterognathiidae; or consist of wings and set with rows of teeth, as in the Gnathostomulidae. The jaws may be solid and forcepslike, as in the order Filospermoidea, or hollow like a pair of forward-pointing funnels, with muscles inserting from behind, as in most Bursovaginoidea. In most species, the inner, anterior parts of the jaw are set with groups or rows of teeth.

Distribution

Gnathostomulids are distributed worldwide, with most species known from the North Atlantic and South Pacific Oceans.

Habitat

Gnathostomulids occur exclusively, sometimes in large numbers, in detritus-rich marine sand as typically found on sheltered beaches, near sea grasses and mangroves, and between coral reefs. Such habitats are often characterized by low oxygen but high hydrogen sulfide concentrations (that create a rotten egg smell), which gnathostomulids seem to tolerate. Most species have been found in the intertidal and shallow subtidal zones, with occasional finds at 1,310 ft (400 m).

Behavior

Gnathostomulids glide through the interstices between sand grains, propelled slowly by their sparse ciliation, and contracting when disturbed. Some species spin a mucous cocoon which may let them survive a deteriorating environment.

Feeding ecology and diet

Belying their fearsome jaws, gnathostomulids are not predators but seem to graze on the microflora (bacteria and fungal threads) attached to sand grains. Scientists do not yet know what role the basal plate and jaws play in feeding.

Reproductive biology

All gnathostomulids are hermaphrodites. The male organs are located in the posterior part of the body. They consist of either an unpaired or paired testes, and a copulatory organ (penis) which, in Scleroperalia, contains a tubelike penis stylet. The sperm is diverse. In Filospermoidea, it is threadlike, with a single ciliary tail of 9+2 microtubules. In Bursovaginoidea, it is aflagellate and droplet shaped, in Conophoralia, aflagellate and cone shaped. Sperm is transferred by copulation, and stored either freely between gut and epidermis, or in a storage pouch (the bursa copulatrix). In all species, the single, unpaired ovary is located dorsally, behind the mouth. Only one large egg matures at a time, and is presumably fertilized by sperm from the bursa before being laid via rupture of the dorsal body wall. Development is direct, and cleavage probably follows the spiral pattern.

Conservation status

No species of gnathostomulid is listed by the IUCN.

Significance to humans

None known.

Species accounts

List of Species

Red haplognathia
Gnathostomula paradoxa
Austrognathia australiensis

Red haplognathia

Haplognathia ruberrima

family

Haplognathiidae

taxonomy

Haplognathia ruberrima Sterrer, 1966, Swedish west coast.

other common names

None known.

physical characteristics

Length to 0.138 in (3,500 µm); diameter 0.0051 in (140 µm); one of the largest gnathostomulids. Most specimens are uniformly brick red, reddish brown, or pink, owing to pigment granules in the epidermis. Rostrum (head) slender and pointed, without paired sensory bristles; posterior end uniformly rounded. Jaws solid, with large winglike apophyses and many sharp denticles; basal plate shieldlike, set with dorsal thorns. Sperm threadlike, with corkscrewlike head.

distribution

The most globally distributed species, found in Australia, Fiji, and Hawaii, as well as on both sides of the North Atlantic and the Mediterranean.

habitat

Like most gnathostomulids, prefers detritus-rich sand in shallow sublittoral areas.

behavior

An animal that has been isolated from the sediments usually coils up by muscular action, then uncoils again by means of its cilia, often from both ends simultaneously, with the rostrum pulling forward and the posterior end pulling backward.

feeding ecology and diet

Seems to graze on fungal hyphae and bacteria adhering to sand grains.

reproductive biology

Single egg is laid by rupture of the dorsal body wall. Egg then sticks to a sand grain until a fully ciliated hatchling 330 µm long emerges.

conservation status

Not listed by the IUCN.

significance to humans

None known.


No common name

Gnathostomula paradoxa

family

Gnathostomulidae

taxonomy

Gnathostomula paradoxa Ax, 1956, Kieler Bucht (Kiel Bay), North Sea.

other common names

None known.

physical characteristics

Length to 0.0394 in (1,000 µm), diameter to 0.00295 in (75 µm). Body colorless; rostrum rounded, headlike, set with paired sensory bristles; posterior drawn out into a short tail. Jaws hollow, with three rows of teeth; basal plate with paired lateral and rostral wings. Reproductive system with a cuticular bursa and a penis stylet. Sperm small, aflagellate, droplet shaped.

distribution

North Sea.

habitat

Occurs in detritus-rich sand, straying into clean sand.

behavior

Nothing is known.

feeding ecology and diet

Grazes on attached microflora.

reproductive biology

Nothing is known.

conservation status

Not listed by the IUCN.

significance to humans

None known.


No common name

Austrognathia australiensis

family

Austrognathiidae

taxonomy

Austrognathia australiensis Sterrer, 2001, Lizard Island, Queens-land, Australia.

other common names

None known.

physical characteristics

Length to 0.0315 in (800 µm), diameter to 0.00315 in (80 µm). Body colorless, slender; head rounded, set with paired sensory bristles; posterior end pointed. Jaws hollow, with two rows of teeth; basal plate with pronounced median and a pair of lateral lobes. Bursa and penis without hard structures; sperm large, cone shaped, aflagellate.

distribution

Queensland, Australia.

habitat

Occurs in detritus-rich sand near sea grass beds and patch reefs.

behavior

Nothing is known.

feeding ecology and diet

Nothing is known.

reproductive biology

Nothing is known.

conservation status

Not listed by the IUCN.

significance to humans

None known.


Resources

Periodicals

Sterrer, W. "Gnathostomulida from the (Sub) tropical Northwestern Atlantic." Studies on the Natural History of the Caribbean Region 74 (1998): 1–178.

——. "Gnathostomulida from Australia and Papua New Guinea." Cahiers de Biologie Marine 42 (2001): 363–395.

Sørensen, M. V., and Sterrer, W. "New Characters in the Gnathostomulid Mouth Parts Revealed by Scanning Electron Microscopy." Journal of Morphology 253 (2002): 310–334.

Wolfgang Sterrer, PhD