Calycophorae

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Calycophorae
Muggiaeaatlanticanectophor.jpg
Morphology of a Calycophorae with aligned nectophores

(Muggiaea atlantica)

Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Cnidaria
Class: Hydrozoa
Order: Siphonophorae
Suborder: Calycophorae
Leuckart, 1854
Families
Hippopodius hippopus with several nectophores. Hippopodius hippopus.jpg
Hippopodius hippopus with several nectophores.
A typical sphaeronectidae with one nectophore (Spheronectes gracilis) FMIB 41696 Sphaeronectes gracilis After Mayer.jpeg
A typical sphaeronectidae with one nectophore (Spheronectes gracilis)

Calycophorae is a suborder of Siphonophores alongside two other suborders Physonectae and Cystonectae. [1] This suborder includes the giant siphonophore, ( Praya dubia ); one of the longest lengthwise extant creatures (40–50m). While the Physonectae have a pneumatophore (a float), nectophore (or nectosome), and a siphosome, Cystonectae lack a nectophore, and Calycophorae lack a pneumatophore. [1] From the bell-shaped nectophores, Physonectae and Calycophorae are called Codonophores or Greek for bell-bearers. [2] The distribution, morphology, and behaviors of Calycophorae species are vast and greatly depend on the species. Calycophoraes typically consist of two nectophores with a siphosome that have many tentacles that grow out of the siphosome. [2] The Calycophoraes move by propelling water out of the nectophore much like how jellyfishes move. The tentacles act as fishing nets where the nematocysts on the tentacles paralyze their prey which are then later fed on. Calycophorae have three life stages, which are the larval development stage, the polygastric stage, and the eudoxid maturation stage. [3] Each Calycophorae colony forms from one fertilized egg. [4]

Distribution

Calycophorae are found generally in every deep sea pelagic environments. [1] [2] [5] The location and depth of distribution depends on the species of Calycophorae. [2] [5] The species Muggiaea bargmannae belongs to the family Diphyidae which have been found at depths between 400 and 2000 meters at both poles at a latitude between 36~87°N and 43~71°S. [2] The species Hippopodius hippopus of the family Hippopodiidae on the other hand, have been observed to live at a depth between 0 and 300 meters at the latitudes 50°N~38°S. [2]

Morphology

All Siphonophores including Calycophorae are composed of zooids, which each have a function within the organism including but not limited to feeding, reproduction, and locomotion. [1] The body plan of a Calycophorae can be divided into its nectophore and its siphosome. [2] A typical Calycophorae has two nectophores that have ostia that are either facing the opposite or same directions. [2] For aligned nectophores, the nectophores are classified as anterior and posterior with the latter being the one behind. [2] Within the nectophore are the somatocyst and the hydroecium. [2] The somatocyst is an additional structure that is seen in the mesoglea of Calycophorans where oil globules are stored as a food source and also serve to provide buoyancy. [2] The hydroecium is a ventral cavity that exists in the posterior nectophore. [2] The siphosome is the stem of the Calycophoran on which the bracts, gastrozooids, and gonophores are attached to with the bracts serving as a protective cover. [1] [2] The bracts also contain a phyllocyst that also contains oil globules. [2] Each group of these zooids is called a cormidium and repeats itself throughout the siphosome. [1] [2] Each gastrozooid has an elongated tentacle that has several side tentacles called tentilla that have pads or batteries of nematocysts which are connected to the tentacle by a pedicel. [2] There are some exceptions to this common morphology such as the Hippopodiids that have several overlapping nectophores and Sphaeronectids that have one nectophore. [2]

Locomotion

The nectophores contract to push water out of the ostium and propels the organism forward. [1] In some Calycophorans, the siphosome can be retracted into the hydroecium to reduce drag. [2] Some Calycophorans have replaced up to 75% of heavy sulfate ions in their bracts with lighter chloride ions to increase their buoyancy to compensate for their lack of pneumatophores. [2]

Feeding

All Siphonophores are predatory carnivores including the Calycophorans. [4] The nematocysts on the tentilla inject toxins into prey that come in contact with the nematocysts. [2] [4] The toxins paralyze the prey immobilizing it, which is then brought to the gastrozooid for digestion and redistribution of nutrients. [4] The prey of Calycophorans depend on the species but are commonly small crustaceans such as copepods, gelatinous zooplankton, or even small fish. [4] [6]

Reproduction

The Calycophorae are monoecious and have three stages in their life cycle which are the larval development stage, the polygastric stage (maturity stage), and the eudoxid stage (sexual reproduction stage). [2] [3] The bracts which contain gastrozooids and gonophores on a Calycophorae detach from the siphosome once maturity is reached. [2] This detached bract becomes a free living eudoxid that releases male or female gametes depending on its determined sex. [2] [7] A successfully fertilized gamete enters the larval development stage in which asexual reproduction takes over to recreate a full polygastric colony. [3] [7]

Taxonomy

From molecular phylogenetics it has been confirmed that Calycophorae evolved from the Physonectae. [1] 175 species of Siphonophores have been identified of which 5 are of Cystonectae (which includes the Portuguese man o' war) and the rest are of Codonophora. [2] There are 16 families under Codonophora and 7 families under Calycophorae: Prayidae, Hippopodiidae, Clausophyidae, Sphaeronectidae, Diphyidae, Abylidae, and Tottonophyidae. [2] [8]

Other relevant information

Current debates

The differences in the structure and function of zooids are often studied to determine the different species of Siphonophores, and the loss or addition of zooids usually signify speciation. [1] Current debates contemplate whether or not the Siphonophores including Calycophorans as a whole are losing or gaining zooid types. [1]

Specimen collection

Siphonophores in general are fragile and using nets would shred the organism into pieces. [1] For descriptive analysis, specimens need to be carefully collected by research submersibles into capsules which takes much more time and effort. [3] These complications are a setback to the study of Calycophorans. [3] However, some locations such as the bay at Villefranche-Sur-Mer, have special abiotic conditions that push up Siphonophores to the shore. [3] These specimens are readily collectible and have contributed to much of our knowledge of Siphonophores. [3]

Related Research Articles

<span class="mw-page-title-main">Cnidaria</span> Aquatic animal phylum having cnydocytes

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.

<span class="mw-page-title-main">Portuguese man o' war</span> Marine invertebrate

The Portuguese man o' war, also known as the man-of-war or bluebottle, is a marine hydrozoan found in the Atlantic Ocean and the Indian Ocean. It is considered to be the same species as the Pacific man o' war or bluebottle, which is found mainly in the Pacific Ocean. The Portuguese man o' war is the only species in the genus Physalia, which in turn is the only genus in the family Physaliidae.

<span class="mw-page-title-main">Hydrozoa</span> Class of cnidarians

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.

<span class="mw-page-title-main">Siphonophorae</span> Order of colonial hydrozoans with differentiated zooids

Siphonophorae 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 species described thus far.

<i>Praya dubia</i> Species of hydrozoan

Praya dubia, the giant siphonophore, lives in the mesopelagic zone to bathypelagic zone at 700 m (2,300 ft) to 1,000 m (3,300 ft) below sea level. It has been found off the coasts around the world, from Iceland in the North Atlantic to Chile in the South Pacific.

<span class="mw-page-title-main">Leptothecata</span> Order of cnidarians with hydrothecae

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 Siphonophorae, which were placed outside the "Hydroida". Given that there are no firm rules for synonymy for high-ranked taxa, alternative names like Leptomedusa, Thecaphora or Thecata, with or without the ending emended to "-ae", are also often used for Leptothecata.

<i>Marrus orthocanna</i> Species of hydrozoan

Marrus orthocanna is a species of pelagic siphonophore, a colonial animal composed of a complex arrangement of zooids, some of which are polyps and some medusae. Swimming independently in the mid-ocean, it lives in the Arctic and other cold, deep waters. It is a colonial creature that is born from a single egg which is fertilized. Later on, a protozoan forms that eventually grows to form more duplicating members of the colony. It belongs to the order Siphonophorae and the genus Marrus, which also includes M. antarcticus, M. claudanielis, and M. orthocannoides.

<i>Apolemia</i> Family of cnidarians

Apolemia is a genus of siphonophores. It is the only genus in the monotypic family Apolemiidae.

<i>Porpita prunella</i> Species of hydrozoan

Porpita prunella is a marine species of hydrozoan organisms within the family Porpitidae. It consists of colonies of zooids. Very little is known about this species, as there have been no confirmed sightings since its discovery in 1801 and naming by Haeckel in 1888. Being in the chondrophore group, it is likely that its behaviour is similar to the other species of the genera in the family. However there are also serious doubts as to its very existence as a separate species and may in fact be a synonym for Porpita porpita instead.

<i>Muggiaea atlantica</i> Species of hydrozoan

Muggiaea atlantica is a species of small hydrozoan, a siphonophore in the family Diphyidae. It is a cosmopolitan species occurring in inshore waters of many of the world's oceans, and it has colonised new areas such as the North Sea and the Adriatic Sea. It is subject to large population swings, and has been held responsible for the death of farmed salmon in Norway. The species was first described by J.T. Cunningham in 1892 from a specimen obtained at Plymouth, England.

Muggiaea kochii is a species of small hydrozoan, a siphonophore in the family Diphyidae.

In biology, Gonozooids are any of the reproductive individuals of Tunicates, Bryozoan, or Hydrozoan colonies that produce gametes. Gonozooids may play a role in labour division or in alternation of generations. A gonozooid typically has hardly any other function than reproduction, amounting to little more than a motile gonad.

<i>Bathyphysa conifera</i> Species of siphonophore sometimes called the flying spaghetti monster

Bathyphysa conifera, sometimes called the flying spaghetti monster, is a bathypelagic species of siphonophore in the family Rhizophysidae. It is found in the northern Atlantic Ocean and off the coast of Southwestern Africa and California.

<span class="mw-page-title-main">Physonectae</span> Suborder of siphonophores

Physonectae is a suborder of siphonophores. In Japanese it is called 胞泳.

<i>Bassia bassensis</i> Species of cnidarian

Bassia is a monotypic siphonophore genus in the family Abylidae. The genus contains the single species Bassia bassensis.

Nanomia bijuga, first described by Stefano Delle Chiaje in 1844 and originally named Physsophora bijuga, is a species of mesopelagic siphonophore in the family Agalmatidae As with all members of the siphonophorae order, it is a colonial organism composed of individual zooids. N. bijuga has a fairly broad distribution, and has been observed in the coastal waters off of North America and Europe. The species has been found to occupy both epipelagic and mesopelagic depths. They utilize specialized swimming zooids for both propulsion and escape behaviors. Similar to other siphonophores, Nanomia bijuga employ stinging tentacles for hunting and defense. They primarily feed on small crustaceans, especially krill.

<i>Abylopsis tetragona</i> Species of cnidarian

Abylopsis tetragona is a species of siphonophore in the family Abylidae.

<span class="mw-page-title-main">Zancleidae</span> Family of cnidarians

Zancleidae is a family of cnidarians belonging to the order Anthoathecata.

Lensia is a genus of hydrozoans belonging to the order Siphonoporae and the family Diphyidae. This genus is colonial and consists of many different types of highly specialized zooids. The genus Lensia was first established in 1932 by Dr. Arthur Knyvett Totton, who would also describe and add another 11 species during his career. As of March 2023, the genus consists of only 26 described and accepted species and an additional seven uncertain species, according to the World Register of Marine Species.

Tima nigroannulata, commonly known as the elegant jellyfish, is a recently discovered colonial hydrozoa found on the Pacific coast of Japan.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 Dunn, Casey; Rugh, Philip; Haddock, Steven (2005-12-01). "Molecular Phylogenetics of the Siphonophora (Cnidaria), with Implications for the Evolution of Functional Specialization". Systematic Biology. 54 (6): 916–935. doi:10.1080/10635150500354837. PMID   16338764 – via Oxford Academic.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Mapstone GM (2014-02-06). "Global diversity and review of Siphonophorae (Cnidaria: Hydrozoa)". PLOS ONE. 9 (2): e87737. Bibcode:2014PLoSO...987737M. doi: 10.1371/journal.pone.0087737 . PMC   3916360 . PMID   24516560.
  3. 1 2 3 4 5 6 7 Dunn C (2009). "Siphonophores". Current Biology. 19 (6): R233-4. doi: 10.1016/j.cub.2009.02.009 . PMID   19321136. S2CID   235312042 . Retrieved 2022-04-25.
  4. 1 2 3 4 5 "Pelagic Siphonophore". www.aquariumofpacific.org. Retrieved 2022-04-25.
  5. 1 2 Grossmann MM, Collins AG, Lindsay DJ (2014-04-03). "Description of the eudoxid stages of Lensia havock and Lensia leloupi (Cnidaria: Siphonophora: Calycophorae), with a review of all known Lensia eudoxid bracts". Systematics and Biodiversity. 12 (2): 163–180. doi:10.1080/14772000.2014.902867. ISSN   1477-2000. S2CID   35309849.
  6. ↑ Pagès F, González HE, Ramón M, Sobarzo M, Gili JM (2001-01-26). "Gelatinous zooplankton assemblages associated with water masses in the Humboldt Current System, and potential predatory impact by Bassia bassensis (Siphonophora: Calycophorae)". Marine Ecology Progress Series. 210: 13–24. Bibcode:2001MEPS..210...13P. doi: 10.3354/meps210013 . hdl: 10261/184183 . ISSN   0171-8630.
  7. 1 2 Carré C, Carré D, Bone Q (1991-10-29). "A complete life cycle of the calycophoran siphonophore Muggiaea kochi (Will) in the laboratory, under different temperature conditions: ecological implications". Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences. 334 (1269): 27–32. Bibcode:1991RSPTB.334...27C. doi:10.1098/rstb.1991.0095.
  8. ↑ Schuchert P (2010-01-06). "WoRMS - World Register of Marine Species - Calycophorae". www.marinespecies.org. Retrieved 2022-04-26.
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