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Dicyemida, also known as Rhombozoa, is a phylum of tiny
parasite 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 h ...
s that live in the renal appendages of
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s.


Taxonomy

Classification is controversial. Traditionally, dicyemids have been grouped with the Orthonectida in the Mesozoa, and, as of 2017, molecular evidence appears to confirm this. However, other molecular phylogenies have placed the dicyemids more closely related to the
roundworms 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 ...
. Additional molecular evidence suggests that this phylum is derived from 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, bra ...
. The phylum is not divided in classes or
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
, but contains three families, Conocyemidae,
Dicyemidae The Dicyemidae is a family of tiny parasites that live in the renal appendages of cephalopods. It contains the following genera and species: * ''Dicyema'' von Kolliker, 1849 * '' Dicyemennea'' Whitman, 1883 * '' Dicyemodeca'' ** '' Dicyemodeca ...
, and Kantharellidae.


Anatomy

Adult dicyemids range in length from , and they can be easily viewed through a light microscope. They display eutely, a condition in which each adult individual of a given species has the same number of cells, making cell number a useful identifying character. Dicyemida lack respiratory, circulatory, excretory, digestive, and nervous systems. The organism's structure is simple: a single axial cell is surrounded by a jacket of twenty to thirty
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proje ...
ted cells. The anterior region of the organism is termed a ''calotte'' and functions to attach the parasite to folds on the surface of its host's renal appendages.


Life cycle

Dicyemids exist in both asexual and sexual forms. The former predominate in juvenile and immature hosts, and the latter in mature hosts. The asexual stage is termed a ''nematogen''; it produces ''vermiform''
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. ...
e within the axial cell. These mature through direct development to form more nematogens. Nematogens proliferate in young cephalopods, filling the kidneys. As the infection ages, perhaps as the nematogens reach a certain density, vermiform larvae mature to form ''rhombogens'', the sexual life stage, rather than more nematogens. This sort of density-responsive reproductive cycle is reminiscent of the asexual reproduction of sporocysts or rediae in larval
trematode Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive h ...
infections of
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
s. As with the trematode asexual stages, a few nematogens can usually be found in older hosts. Their function may be to increase the population of the parasite to keep up with the growth of the host. Rhombogens contain hermaphroditic
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 sp ...
s developed within the axial cell. These gonads, more correctly termed ''infusorigens'', self-fertilise to produce ''infusoriform'' larvae. These larvae possess a very distinctive morphology, swimming about with ciliated rings that resemble headlights. It has long been assumed that this sexually produced infusoriform, which is released when the host eliminates urine from the kidneys, is both the dispersal and the infectious stage. The mechanism of infection, however, remains unknown, as are the effects, if any, of dicyemids on their hosts. Some part of the dicyemid life cycle may be tied to
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
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 " ...
environments, where they occur in greatest abundance. While dicyemids have occasionally been found in the tropics, the infection rates are typically quite low, in and many potential host species are not infected. Dicyemids have never been reported from truly oceanic cephalopods, who instead host a parasitic
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...
fauna. Most dicyemid species are recovered from only one or two host species. While not strictly host specific, most dicyemids are only found in a few closely related hosts.


References


Further reading

* * * * * {{Taxonbar, from1=Q134869, from2=Q21447146, from3=Q2224427 Animal phyla Parasitic protostomes Parasites of molluscs