Euplokamis
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''Euplokamis'' is a genus of
ctenophore Ctenophora (; : ctenophore ) is a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and they ar ...
s, or comb jellies, belonging to the monotypic family Euplokamididae. It shares the common name sea gooseberry with species of the genus Pleurobrachia. After being originally described by Chun (1879), the family '' Euplokamididae'' was expanded by Mills (1987) due to the discovery of a new species, '' Euplokamis dunlapae''. Further research indicated that ''Euplokamis'' should be identified from Mertensiidae due to the rows of combs and some compression. They may also be distinguished from the genus '' Pleurobrachia'' due to their more elongated shape. Additionally, various adaptations of ''Euplokamis'' have been observed such as the use of tentacles for movement/feeding, a complex nervous system, and bioluminescent capabilities. Other characteristics including a defined
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical ...
, lack of stinging cells, developmental differences, and symmetry supported the reclassification of these organisms.


Distribution & Habitat

''Euplokamis'' have been found in fossil records dating back to the
Cambrian period The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordovici ...
, part of the
Paleozoic Era The Paleozoic ( , , ; or Palaeozoic) Era is the first of three geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma at the start of ...
, and it is estimated that some species may have evolved before this period. Originally,
Cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
ns and
Ctenophores Ctenophora (; : ctenophore ) is a phylum of marine invertebrates, commonly known as comb jellies, that marine habitats, inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs ...
were classified under the same phyla,
Coelenterata Coelenterata is a term encompassing the animal phyla Cnidaria (corals, true jellies, sea anemones, sea pens, and their relatives) and Ctenophora (comb jellies). The name comes , referring to the hollow body cavity common to these two phyla. Th ...
. Ctenophore bodies are made up of a gelatin substance, similar to
Cnidarians Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
, but the multiple rows of combs present in fossil records are unique to ctenophores. Records of ''Euplokamis'' sp. indicate they are distributed widely around the world, but are most often found in warm coastal waters. ''Euplokamis'' prefer marine, or saltwater, environments and are classified as free swimmers, due to their ability to move through the water column. They have been identified in the Mediterranean Sea, the North Pacific, the Gulf of Maine, and off the coast of Sweden. However, since this genus was originally grouped in the family Pleurobrachiidae, there is limited information regarding their actual distribution and habitat.


Anatomy & Morphology

Ctenophores are divided into two classes based on either the presence(
Tentaculata Tentaculata is a class of comb jellies, one of two classes in the phylum Ctenophora. The common feature of this class is a pair of long, feathery, contractile tentacles, which can be retracted into specialised ciliated sheaths. In some species, ...
)) or the lack (
Nuda Beroidae is a family of ctenophores or comb jellies more commonly referred to as the beroids. It is the only known family within the monotypic order Beroida and the class Nuda. They are distinguished from other comb jellies by the complete abse ...
) of tentacles. Within each class, there are multiple orders to further distinguish their structures and characteristics. The class
Tentaculata Tentaculata is a class of comb jellies, one of two classes in the phylum Ctenophora. The common feature of this class is a pair of long, feathery, contractile tentacles, which can be retracted into specialised ciliated sheaths. In some species, ...
contains the following orders:
Cydippida Cydippida is an order of comb jellies. They are distinguished from other comb jellies by their spherical or oval bodies, and the fact their tentacles are branched, and can be retracted into pouches on either side of the pharynx. The order is not ...
,
Lobata Lobata is an order of transparent marine invertebrates belonging to the phylum of Ctenophora in the class Tentaculata, and are commonly referred to as comb jellies or sea gooseberries. There are currently 19 extant known species in the order of ...
, and
Cestida Cestidae is a family of comb jellies. It is the only family in the monotypic order Cestida. Unlike other comb jellies, the body of cestids is greatly flattened, and drawn out into a long ribbon-like shape. The two tentacles are greatly shortened, ...
. The genus ''Euplokamis'' is part of the class tentaculate, which indicates that tentacles are present. They are also part of the order
Cydippida Cydippida is an order of comb jellies. They are distinguished from other comb jellies by their spherical or oval bodies, and the fact their tentacles are branched, and can be retracted into pouches on either side of the pharynx. The order is not ...
, distinguished by their tentacles and their round body shape. ''Euplokamis'' tentacles are long, with side branches, and have a sheath allowing them to be retracted inside of the body. The tentacle side branches are known as tentilla, which in the case of ''Euplokamis'' are held tightly in coils except during the act of prey capture. Further, the widely spaced tentillia droplets allow for organisms to be classified to the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
level and are one of the only examples of striated muscle found in ctenophores. Additionally, these organisms have bi-radial symmetry with a mouth on their front end and a
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, crustaceans, and gastropods, A similar structure is also found in '' Xenoturbella''. T ...
, or sense organ, at the other end. The sides of their stomachs are lined with distinct bulbs, shaped like tadpoles, and unlike other well-known jellyfish, ''Euplokamis'' do not have any nematocytes, known as stinging cells. The mouth is connected to the digestive tract via the
pharynx The pharynx (: pharynges) is the part of the throat behind the human mouth, mouth and nasal cavity, and above the esophagus and trachea (the tubes going down to the stomach and the lungs respectively). It is found in vertebrates and invertebrates ...
. The digestive system, or
gastrovascular cavity The gastrovascular cavity is the primary organ of digestion and circulation in two major animal phyla: the Coelenterates or cnidarians (including jellyfish and corals) and Platyhelminthes (flatworms). The cavity may be extensively branched into ...
, is made up of intricate canals that allow for both digestion and circulation to occur. They also lack an
anus In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
but are able to excrete some waste through pores on the adoral end. Typically, ''Euplokamis'' are small, only growing up to approximately 20 millimeters (mm) in length. Another distinct feature of this genus is the eight rows of combs present. While they are known as comb plates, they are actually made up of large cilia, which are hair-like structures. These plates are unique because they consist of some of the largest known cilia found on any organism. Additionally, the combs primarily function in movement, allowing some species to move forward and backward. Since their bodies are made of the
mesoglea Mesoglea refers to the extracellular matrix found in cnidarians like coral or jellyfish as well as ctenophores that functions as a hydrostatic skeleton. It is related to but distinct from mesohyl, which generally refers to extracellular material f ...
—a translucent, gelatin-like substance—the 8 comb rows can be easily identified. These combs function in movement, due to their large ciliary structures.


Behaviors & Adaptations


Tentacles: Movement and Feeding Behavior

''Euplokamis'' have long tentacles with branches that are used for feeding and movement. These branches are known as tentilla and are held tightly in coils, forming droplet shapes. The tentilla are usually held in coils but can be uncoiled to aid in movement. Further, these organisms can move their tentilla in slow spontaneous movements or in rapid extensions. According to research, ''Euplokamis'' are
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
, like all other known species of Ctenophores. They are known to feed primarily on
rotifer The rotifers (, from Latin 'wheel' and 'bearing'), sometimes called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic Coelom#Pseudocoelomates, pseudocoelomate animals. They were first describ ...
s and small
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s: including
copepod Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (living in the water column), some are benthos, benthic (living on the sedimen ...
s,
amphipods Amphipoda () is an order (biology), order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods () range in size from and are mostly detritivores or scavengers. There are more than 10,700 amphip ...
, and some planktonic larvae. To catch prey, they will extend the tentilla and wrap them around copepods. The tentilla are covered in sticky colloblasts, which keep the prey stuck in place. Not only can the tentilla be released at high velocities to quickly capture prey, but they also can be released in a slow and controlled manner, likely to attract prey. Additionally, the cilia that make up the comb plate move by making a stroke in a certain direction. After capturing prey, they can reverse the stroke, or beat, in the other direction on two rows, while the other rows continue to beat in the normal direction. They are then able to push the prey to their mouth in a sweeping motion, and the ciliary reversal causes the organism to rotate, which tangles the prey further into its mouth.


Nervous System

Despite their simple exterior, research on the nervous system of ''Euplokamis'' sp. indicates the use of more complex systems, including axons. The use of these axons has allowed some species of ''Euplokamis'' sp. to swim backward rapidly. The direction of the cilia comb plate may be reversed causing them to move backward. These organisms are unique due to the presence of giant axons in their combs that allow for a rapid escape response. Additionally, ''Euplokamis'' sp. has an aboral sensory organ, which is bypassed to produce this escape response.


Bioluminescence

Another adaptation that many ctenophores have developed is bioluminescence, or the ability to produce light. For example, ''Euplokamis dunlapae'' were found to produce light off the coast of Washington when exposed to stimuli. After physical stimulation, they produced bright flashes of light, consistent with bioluminescence. According to research, bioluminescence in ''Euplokamis'' sp. is both intrinsic and extrinsic, because light can appear in the comb rows or as bursts of light in the water. Additionally, off the coast of Maine, ''Euplokamis'' sp. was found to be one of the two brightest species to have bioluminescence. Research suggests that bioluminescence in ''Euplokamis'' sp. may function as a defense mechanism. The bursts of light were only observed directly in response to a disturbance or stimulation, likely to distract or blind predators when they are sensed. Additionally, some produce light as a warning signal, or to expose nearby predators. The various strategies may impact predator-prey relationships or other the population dynamics in an area.


Taxonomy

Source: * Family: Euplokamididae (Mills, 1987) ** Genus: ''Euplokamis'' (Chun, 1879) *** Species: '' Euplokamis crinita'' (Moser, 1909) *** Species: '' Euplokamis dunlapae'' (Mills, 1987) *** Species: '' Euplokamis evansae'' (Gershwin, Zeidler & Davie, 2010) *** Species: '' Euplokamis helicoides'' (Ralph & Kaberry, 1950) *** Species: '' Euplokamis octoptera'' (Mertens, 1833) *** Species: '' Euplokamis stationis'' (Chun, 1879) Changes: * Species: ''Euplokamis californiensis'' (Torrey, 1904) accepted as '' Hormiphora californensis'' (Torrey, 1904) * Species: ''Euplokamis cucumis'' (accepted as '' Hormiphora cucumis'' (Mertens, 1833)


References

{{Taxonbar, from1=Q3823001, from2=Q21854349 Tentaculata Ctenophore genera Taxa named by Carl Chun