Polypodium hydriforme | |
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Two specimens of free-living Polypodium | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Cnidaria |
Class: | Polypodiozoa Raikova, 1988 |
Order: | Polypodiidea Poche, 1914 |
Family: | Polypodiidae Poche, 1914 |
Genus: | Polypodium Ussov, 1885 |
Species: | P. hydriforme |
Binomial name | |
Polypodium hydriforme | |
Polypodium is a genus of cnidarians that parasitizes in the eggs of sturgeon and similar fishes (Acipenseridae and Polyodontidae). [2] It is one of few animals that lives inside the cells of other animals.
Polypodium hydriforme is the only species of this monotypic genus. The parent family (Polypodiidae), order (Polypodiidea) and class (Polypodiozoa) are also monotypic.
Unusual characteristics have led to much controversy regarding the phylogenetic position of Polypodium within metazoans. [2]
Polypodium has traditionally been considered a cnidarian because it possesses nematocysts, the stinging structures characteristic of this phylum. [2] Molecular phylogenetic studies using 18S rDNA sequence data temporarily challenged this interpretation, by finding that Polypodium is a close relative to myxozoans and suggesting that together they share a closer affinity to bilaterians than cnidarians. [3] Due to the variable rates of 18S rDNA sequences, these results were however suggested to be an artifact of long branch attraction, [2] and myxozoans have in the meantime, also been classified within cnidarians. [4]
Evans et al. (2008) [2] performed phylogenetic analyses of metazoans with 18S and partial 28S rDNA sequences in a large dataset that includes Polypodium and a comprehensive sampling of cnidarian taxa. This supports the placement of Polypodium within Cnidaria. [2] This accords with its traditional classification, in particular with the fact that Polypodium possesses nematocysts and a cnidarian-like body plan. [2] Myxozoans are currently classified as cnidarians as well. [4]
Polypodium was discovered in 1871 by Professor Owsiannikov within the eggs of sterlet (Acipenser ruthenus) from the Volga river in Russia. In 1885 Ussov named Owsiannikov's "parasitic larva" Polypodium hydriforme and gave a morphological description of the parasite. [5] Polypodium was long considered a unique intracellular parasite among cnidarians. [6] [7] Its hosts include 14 species of Acipenser , 2 species of Huso , Polyodon spathula [6] and Scaphirhynchus platorynchus . [2]
Polypodium hydriforme is an endocellular parasite with an unusual life cycle, a peculiar morphology, and high rates of DNA evolution. Polypodium spends most of its life inside the oocytes of acipenseriform fishes. In infected oocytes, Polypodium develops from a binucleate cell into an inside-out planuliform larva and then into an elongate inside-out stolon; the epidermal cell layer is located internal to the body and the gastrodermis is located externally. [2] [6] [8] The embryo, larva and stolon are surrounded by a protective polyploid cell, which also functions in digestion. [6] Just prior to host spawning, Polypodium everts to the normal position of cell layers, revealing tentacles scattered along the stolon. [2] During eversion, the yolk of the host oocyte fills the gastral cavities of the parasite, supplying the future free-living stage with nutrients. [2] [8] The parasitic phase of its life cycle usually takes several years. Finally, upon emerging from the host egg in fresh water, the free-living stolon fragments into individual medusoid-like organisms that go on to multiply by means of longitudinal fission. In summer they form endodermal sexual organs: "female" ones showing ovaria and gonoducts, and "male" ones with simpler organization. [8] "Female" gonads are supposedly abortive; [6] [8] the "male" ones ultimately produce binucleate cells [6] [8] and become gametophores which infect host fish. [6] Polypodium hydriforme displays many peculiar characteristics, some of them shared with myxozoa. [2] [7]
The habitat of Polypodium hydriforme is freshwater. [2]
Although freshwater is an unusual habitat for cnidarians, it is not unheard of, especially within hydrozoans. [2] For instance, the model organism Hydra and the jellyfish Craspedacusta sowerbii are both exclusively freshwater hydrozoans. [2] Hydra and Craspedacusta sowerbii are distantly related and are not closely related to Polypodium. Also, the obligate parasite Myxobolus cerebralis lives in freshwater. Thus, it appears that in the evolution of cnidarians, invasion of freshwater habitats has happened at least three separate times. [2]
Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in fresh water and marine environments, including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are a decentralized nervous system distributed throughout a gelatinous body and the presence of cnidocytes or cnidoblasts, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living, jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians are also some of the only animals that can reproduce both sexually and asexually.
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm of ciliated cells surrounding one or more reproductive cells.
Myxozoa is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. Over 2,180 species have been described and some estimates have suggested at least 30,000 undiscovered species. Many have a two-host lifecycle, involving a fish and an annelid worm or a bryozoan. The average size of a myxosporean spore usually ranges from 10 μm to 20 μm, whereas that of a malacosporean spore can be up to 2 mm. Myxozoans can live in both freshwater and marine habitats.
Hydrozoa is a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specialized individual animals cannot survive outside the colony. A few genera within this class live in freshwater habitats. Hydrozoans are related to jellyfish and corals and belong to the phylum Cnidaria.
Myxosporea is a class of microscopic animals, all of whom are parasites. They belong to the Myxozoa clade within Cnidaria. They have a complex life cycle that comprises vegetative forms in two hosts—one an aquatic invertebrate and the other an ectothermic vertebrate, usually a fish. Each parasitized host releases a different type of spore. The two forms of spore are so different that until relatively recently they were treated as belonging to different classes within the Myxozoa.
Velella is a monospecific genus of hydrozoa in the Porpitidae family. Its only known species is Velella velella, a cosmopolitan free-floating hydrozoan that lives on the surface of the open ocean. It is commonly known by the names sea raft, by-the-wind sailor, purple sail, little sail, or simply Velella.
Myxobolus cerebralis is a myxosporean parasite of salmonids that causes whirling disease in farmed salmon and trout and also in wild fish populations. It was first described in rainbow trout in Germany in 1893, but its range has spread and it has appeared in most of Europe, the United States, South Africa, Canada and other countries from shipments of cultured and wild fish. In the 1980s, M. cerebralis was found to require a tubificid oligochaete to complete its life cycle. The parasite infects its hosts with its cells after piercing them with polar filaments ejected from nematocyst-like capsules. This infects the cartilage and possibly the nervous tissue of salmonids, causing a potentially lethal infection in which the host develops a black tail, spinal deformities, and possibly more deformities in the anterior part of the fish.
Tetracapsuloides bryosalmonae is a myxozoan parasite of salmonid fish. It is the only species currently recognized in the monotypic genus Tetracapsuloides. It is the cause of proliferative kidney disease (PKD), one of the most serious parasitic diseases of salmonid populations in Europe and North America that can result in losses of up to 90% in infected populations.
A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores, which are not related to cnidarians at all. Some groups of Nemerteans also produce larvae that are very similar to the planula, which are called planuliform larva. In a few cnidarian clades, like Aplanulata and the parasitic Myxozoa, the planula larval stage has been lost.
Medusozoa is a clade in the phylum Cnidaria, and is often considered a subphylum. It includes the classes Hydrozoa, Scyphozoa, Staurozoa and Cubozoa, and possibly the parasitic Polypodiozoa. Medusozoans are distinguished by having a medusa stage in their often complex life cycle, a medusa typically being an umbrella-shaped body with stinging tentacles around the edge. With the exception of some Hydrozoa, all are called jellyfish in their free-swimming medusa phase.
Craspedacusta sowerbii or peach blossom jellyfish is a species of freshwater hydrozoan jellyfish, or hydromedusa cnidarian. Hydromedusan jellyfish differ from scyphozoan jellyfish because they have a muscular, shelf-like structure called a velum on the ventral surface, attached to the bell margin. Originally from the Yangtze basin in China, C. sowerbii is an introduced species now found throughout the world in bodies of fresh water.
Turritopsis dohrnii, also known as the immortal jellyfish, is a species of small, biologically immortal jellyfish found worldwide in temperate to tropic waters. It is one of the few known cases of animals capable of reverting completely to a sexually immature, colonial stage after having reached sexual maturity as a solitary individual. Others include the jellyfish Laodicea undulata and species of the genus Aurelia.
Buddenbrockia plumatellae is a worm-like parasite of bryozoans whose taxonomic placement long puzzled biologists. It is now classified as one of only three myxozoans in the Malacosporea subclass and its only family, Saccosporidae, on the basis of both genetic and ultrastructural studies. It was the first multicellular myxozoan identified and its vermiform shape initially gave strong support to the theory that the enigmatic group belongs among the Bilateria. Five years later, this was refuted by a study of fifty genes from this same "worm", which had rarely even been seen since its discovery in 1851. These 50 phylogenetic markers reveal that Buddenbrockia is closely related to jellyfish and sea anemones, typical members of the animalian phylum Cnidaria. Because of the highly divergent nuclear protein sequences of Buddenbrockia, relative to those of the other animals compared in this study, only the use of a sophisticated tree-building approach allowed for recovery of its cnidarian evolutionary affinities. One of the researchers talked about the problems encountered studying its morphology: “It has no mouth, no gut, no brain and no nerve cord. It doesn’t have a left or right side or a top or bottom – we can’t even tell which end is the front!” As the myxozoans are now demonstrably non-bilaterian in origin, he concluded that “the worm-like body shape evolved at least twice from two completely different kinds of animal.”
Hydroidolina is a subclass of Hydrozoa and makes up 90% of the class. Controversy surrounds who the sister groups of Hydroidolina are, but research has shown that three orders remain consistent as direct relatives: Siphonophorae, Anthoathecata, and Leptothecata.
Paratomy is a form of asexual reproduction in animals where the organism splits in a plane perpendicular to the antero-posterior axis and the split is preceded by the "pregeneration" of the anterior structures in the posterior portion. The developing organisms have their body axis aligned, i.e., they develop in a head to tail fashion.
Clytia hemisphaerica is a small hydrozoan-group cnidarian, about 1 cm in diameter, that is found in the Mediterranean Sea and the North-East Atlantic Ocean. Clytia has the free-swimming jellyfish form typical of the Hydrozoa, as well as vegetatively propagating polyps.
Clava is a monotypic genus of hydrozoans in the family Hydractiniidae. It contains only one accepted species, Clava multicornis. Other names synonymous with Clava multicornis include Clava cornea, Clava diffusa, Clava leptostyla, Clava nodosa, Clava parasitica, Clava squamata, Coryne squamata, Hydra multicornis, and Hydra squamata. The larvae form of the species has a well developed nervous system compared to its small size. The adult form is also advanced due to its ability to stay dormant during unfavorable periods.
Sphaerospora molnari is a microscopic endoparasite of carp in pond cultures and natural freshwater habitats in Central and Eastern Europe. In natural infections, S. molnari invades the epithelia of gills and surrounding skin regions. It then forms spores in between epithelial cells, causing sphaerosporosis, a pathological condition of the skin and gill tissues. Affected tissues show marked dystrophic changes and necrosis, causing secondary bacterial infections and resulting in osmoregulatory and respiratory failure. Mortalities can reach 100% but little is known about the overall distribution of the parasite species in European carp ponds or its economic impact on carp aquaculture.
Thelohanellus kitauei is a myxozoan endoparasite identified as the agent of intestinal giant-cystic disease (IGCD) of common carp Cyprinus carpio. The species was first identified in Japan, in 1980 and later formally described by Egusa & Nakajima. Fan subsequently reported the parasite in China, and several other reports from carp and Koi carp in China and Korea followed. Reports referred to an intestinal infection, swelling and emaciation of fish due to blockage of the intestinal tract by giant cysts. The intestine of carp was believed to be the only infection site of T. kitauei until Zhai et al. reported large cysts of T. kitauei in the skin, with morphologically similar and molecularly identical spores. T. kitauei has been recognized as the most detrimental disease of farmed carp in Asia with around 20% of farmed carp killed annually. In 2014, the genome of T. kitauei was sequenced, and in 2016, its life cycle was found to include the oligochaete Branchiura sowerbyi. Infected oligochaete worms were first discovered in Hungary and raised concerns of the introduction of T. kitauei into European carp culture ponds, since it was believed to be endemic to Asia. However, the related disease (IGCD) has not yet been reported in Europe.
Zancleidae is a family of cnidarians belonging to the order Anthoathecata.
This article incorporates CC-BY-2.0 text from the reference. [2]