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Hydroidolina is a subclass of
Hydrozoa Hydrozoa (hydrozoans; ) are 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 specializ ...
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 Siphonophorae (from Greek ''siphōn'' 'tube' + ''pherein'' 'to bear') is an order within Hydrozoa, which is a class of marine organisms within the phylum Cnidaria. According to the World Register of Marine Species, the order contains 175 specie ...
,
Anthoathecata Anthoathecata, or the athecate hydroids, are an order of hydrozoans belonging to the phylum Cnidaria. A profusion of alternate scientific names exists for this long-known, heavily discussed, and spectacular group. It has also been called Gymnobl ...
, and
Leptothecata Leptothecata, or thecate hydroids, are an order of hydrozoans in the phylum Cnidaria. Their closest living relatives are the athecate hydroids, which are similar enough to have always been considered closely related, and the very apomorphic Sipho ...
.


Description and background

The phylum
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
contains two clades:
Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, Scleractinia, stony corals and Alcyonacea, soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plank ...
and
Medusozoa 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 sta ...
. There are around 3800 species within the clade
Medusozoa 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 sta ...
and it consists of
Cubozoan Box jellyfish (class Cubozoa) are cnidarian invertebrates distinguished by their box-like (i.e. cube-shaped) body. Some species of box jellyfish produce potent venom delivered by contact with their tentacles. Stings from some species, including '' ...
s, Scyphozoans, and
Hydrozoa Hydrozoa (hydrozoans; ) are 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 specializ ...
ns. Hydroidolina are small predatory animals, ranging in 8-30 millimeters in size, exhibiting radial symmetry and are
diploblastic Diploblasty is a condition of the blastula in which there are two primary germ layers: the ectoderm and endoderm. Diploblastic organisms are organisms which develop from such a blastula, and include cnidaria and ctenophora Ctenophora (; ...
(developed from two embryonic layers: ectoderm and endoderm). The classification below is based on the
World Register of Marine Species The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive list of names of marine organisms. Content The content of the registry is edited and maintained by scientific specialist ...
: Subclass Hydroidolina * Order
Anthoathecata Anthoathecata, or the athecate hydroids, are an order of hydrozoans belonging to the phylum Cnidaria. A profusion of alternate scientific names exists for this long-known, heavily discussed, and spectacular group. It has also been called Gymnobl ...
* Order
Leptothecata Leptothecata, or thecate hydroids, are an order of hydrozoans in the phylum Cnidaria. Their closest living relatives are the athecate hydroids, which are similar enough to have always been considered closely related, and the very apomorphic Sipho ...
* Order
Siphonophorae Siphonophorae (from Greek ''siphōn'' 'tube' + ''pherein'' 'to bear') is an order within Hydrozoa, which is a class of marine organisms within the phylum Cnidaria. According to the World Register of Marine Species, the order contains 175 specie ...


Distribution

Hydroidolina are commonly found in a variety of marine environments across the world such as deepwater caves or brackish and fresh shallow waters, and can exist as solitary or colonial. Benthic polyps can be found on a variety of hard substrates, including both natural and artificial surfaces. Many of them live on other organisms such as fish,
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ca ...
s, algae, and
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s. Furthermore, they prefer not to settle on sand or similarly textured surfaces unless fauna or flora is present. Because Hydroidomedusian polyps often settle on other organisms, they are also subject to partake in symbiotic relationships. For example, the bivalve mantle cavity of a
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
provides a sheltered environment, transporting food due to the current. In exchange, the hydroid protects against intruders.


Diet

Hydroidolina are carnivorous suspension feeders. Motile medusa use their
cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this ce ...
s and tentacles to capture prey.


Anatomy and morphology

Cnidarians are united by the common characteristic of having a specialized cell called a cnidocyte, which contains an explosive organelle called a
cnidocyst A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this ce ...
, or stinging cell. In Hydrozoans, the cnidocysts are formed from interstitial stem cells in the ectoderm and are used for prey capture and anti-predator defense.
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
ns are known to occur in two body forms: the polyp form which is benthic and “stalk-like,” and the medusae form, which is commonly known as the “bell” form. Polyp forms are sessile as adults, with a single opening (the mouth/anus) to the digestive cavity facing up with tentacles surrounding it. Medusa forms are motile, with the mouth and tentacles hanging down from an umbrella-shaped bell. Though some outlier
Hydrozoa Hydrozoa (hydrozoans; ) are 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 specializ ...
ns go through a polyploid (polyp) and medusa stage, Hydroidolina, which comprises almost all hydrozoans, goes through an asexual polypoid stage where the polyp fixed to a substrate and a sexual hydroid stage varying from free-swimming medusa to a
gonophore A gonophore is a reproductive organ in Hydrozoa that produces gametes. It is a sporosac, a medusa or any intermediate stage. The name is derived from the Greek words (, that which produces seed) and (, -bearing). Gonophores are borne on branch ...
that remains attached to the polyp. An important characteristic of the Hydroidolina is the presence and formation of an exoskeleton. The exoskeleton varies in chemical composition, structural rigidity, thickness, and coverage within the different regions of the colony and protects the coenosarc of the polypoid stage. It originates as epidermal secretions, with the exosarc being produced first by glandular epidermal cells. The exoskeleton can either be bilayered and contain both the exosarc (outer layer) and perisarc (inner layer) or corneous (just perisarc). The exoskeleton contains anchoring structures such as desmocytes and "perisarc extensions."


Life cycle and reproduction

The Hydroidolina follows a biphasic life cycle, which alternates in occurrence as planula larva, asexual colonial sessile polyps and free-swimming sexual medusa, not all of which may be present in the one life cycle of the Hydroidolina. Within its benthic phase, polyps of these hydroids attach to soft tissues on organisms, such as the mantle of a mollusk, and reproduce asexually by budding In the sexual medusa stage, gonophores, which are the reproductive organ that produces gametes, and will stay attached to the polyp as a reduced medusa stage but will sometimes, often rarely, form to become their own medusae.


Taxonomy


Alternate classifications

Other hydrozoan classifications, which are beset by
paraphyly In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
however, are still often seen. They do not unite the Leptolinae in a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
and thus do not have any merit according to modern understanding of hydrozoan
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
. The alternate name Leptolinae (or Leptolina) was used in older sources for Hydroidolina. The obsolete name Hydroida was used for a paraphyletic grouping that is now considered synonymous with Hydroidolina but did not include the colonial jellies of the order Siphonophorae.


Ecological Impact

The complexity of fauna environments in shallow and deep waters is only increased by benthic polyp colonization. These hydroid colonies affect many spatial and temporal settlement patterns of other benthic species due to providing a habitat for a wide variety of organisms, thus promoting species richness and abundance. These sessile invertebrates could prove to be useful as a measure of environmental changes within their own colonies as well as for changes within near marine environments pertaining to temporal and spatial changes to species distribution and composition, temperature, and food.


References

{{Taxonbar, from=Q1942461 Hydrozoa