Box jellyfish

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Box jellyfish
Carybdea marsupialis ("Mediterranean sea wasp")

Carybdea marsupialis ("Mediterranean sea wasp")

Systematics
without rank: Opisthokonta
without rank: Holozoa
without rank: Multicellular animals (Metazoa)
without rank: Tissue animals (Eumetazoa)
Trunk : Cnidarians (Cnidaria)
Class : Box jellyfish
Scientific name
Cubozoa
Werner , 1975

The box jellyfish (Cubozoa) form a small, almost exclusively marine class of cnidarians (Cnidaria) with currently about 50 species described. The name comes from the cube (to cuboid) shaped umbrella of the medusa. They are solitary cnidarians with a fixed polyp stage and a free- living medusa stage . Medusa formation usually takes place through complete metamorphosis of the polyp, less often through a special form of strobilation (constriction). They occur worldwide mostly in tropical and subtropical seas; the development of polyps also takes place in some species in estuarine areas with changing salt contents. Individual species rarely penetrate into moderate latitudes. Box jellyfish are active, predatory animals whose movement and migration behavior can be compared to fish. They belong to the necton of the seas.

The box jellyfish include the most feared species of jellyfish, including the Carukia barnesi , a species from the "Irukandji" group, which, along with other species , can trigger Irukandji syndrome , and Chironex fleckeri from the group of "sea wasps", whose poison is can kill a person in a matter of minutes.

The oldest fossil box jellyfish comes from the Pennsylvania (carbon). In terms of tribal history, their closest relatives are probably the English jellyfish (Staurozoa).

features

Box jellyfish are metagenetic solitary cnidarians with a comparatively small sessile polyp and a significantly larger free-living medusa in the adult stage . In most species, medusa formation takes place through a complete metamorphosis of the polyp, i.e. the polyp is completely transformed into a single medusa. Recently, however, a second type of medusa formation has been observed in the species Carybdea marsupialis , a modified strobilation in which a regenerative remnant of the polyp remains, which can grow back into a polyp. The polyps are quite small and only measure a few millimeters. The medusa size in the adult stage ranges from about 1 centimeter to about 30 centimeters in height or diameter. Usually the height is significantly greater than the diameter. Some species can develop tentacles up to three meters long , so the size of the umbrella does not say much about the overall size. Screen jelly and all organs of the medusa are transparent. The life cycle of most species is still unknown. Only Carybdea marsupialis and Tripedalia cystophora have been well studied , so that most observations on polyps are limited to these two species.

polyp

The polyp is radially symmetrical without a solid skeleton and usually attached to the ground with a basal disk . In Carybdea marsupialis it is mostly naked, only in Tripedalia cystophora has a peridermal covering in the lower part of the polyp been detected. The surface of Carybdea marsupialis is covered with flagella , in Tripedalia cystophora these are stiff and act as sensory hairs. The body wall consists of the epidermis and gastrodermis with the primarily cell-free mesogloea in between , which acts as a supporting lamella. It has a uniform, sac-shaped gastric space with a nerve ring. In the resting state, the gastrodermis can be folded lengthways. However, these are not partitions (septa), such as B. in the flower animals (Anthozoa), because the folds disappear again when the gastric space is stretched. The tentacles are arranged in a wreath around the mouth; the mouth cone ( proboscis ) with the mouth clearly protrudes above the tentacle rim. It is very elastic and stretchable and can therefore look more like a trunk when stretched. The body area under the tentacle rim is somewhat expanded and contains the stomach. The lower area of ​​the polyp is again somewhat narrower (stem) and ends in an adhesive disk with which the polyp is attached to the substrate. The solid tentacles are only slightly thickened at the base, otherwise they are almost equally thin when extended. When contracted, they are in the shape of a spherical bud, or they can be almost completely withdrawn into the body. In the few species examined, there is usually only a single nettle cell on the slightly thickened tip of each tentacle. Used nettle cells are replaced by migrating nettle cells. In Tripedalia cystophora , on the other hand, there are around 20 to 40 nettle cells in the area of ​​the tip of the tentacle. The polyps have only one type of nettle cells, so-called stenotles . The species examined so far are characterized by a relatively small polyp, which is usually only a few millimeters in size before it metagenetically transforms into a medusa. Even in species whose medusa stage can reach a considerable size (approx. 30 cm), the polyp is only a few millimeters in size. The polyps grow up after about three months. The polyp of Tripedalia cystophora is capable of limited locomotion. It can detach itself from the subsurface and crawl to find a new habitat or substrate. In the creeping stage, it resembles a secondary polyp that has formed by budding on an adult polyp. If living conditions are unfavorable, the polyp can also encapsulate (encyst) itself. It retracts all tentacles, becomes spherical and surrounds itself with a thin peridermal cover. In this state it can also easily be carried by currents. If the living conditions become more favorable again, it detaches itself from the shell and crawls to find a new location where it can attach itself.

Metamorphosis begins with the formation of four longitudinal furrows that divide the top of the polyp into four quadrants. The tentacles, randomly distributed in a circle in polyps, are combined into four groups; the tentacle bases merge. The tentacles are reabsorbed and the four fused tentacle bases are transformed into the rhopals . New tentacles grow in the interradii. The previously uniform gastric space of the polyp is converted into the structured gastric space of the medusa. The manubrium and velarium, a muscular fold of skin that narrows the screen opening, are formed. The basal part is now almost completely included in the upper part of the medusa umbrella. After the musculature of the medusa has differentiated, the basal adhesive disc also recedes and the medusa swims away. As a rule, only a mucous stain remains or, in the case of Tripedalia cystophora, the peridermal cup . The metamorphosis takes about five to seven days at 25 to 27 ° C.

Medusa

The medusa is tetraradial, i.e. That is, it has an approximately square, whole-rimmed umbrella when viewed from above. The edges of the umbrella are rounded, and the height is usually significantly larger than the diameter, so that the umbrella is less often cube-shaped (name!), But more often cuboid. If the lateral edges of the cuboid are strongly rounded, the shape can also be called bell-shaped. The four (rounded) edges of the screen, also called pedal shoulders, are located in the so-called inter-radii, the per-radii form the sides of the cube. The top of the umbrella is curved and is also known as the exumbrella, the inwardly curved underside is called the subumbrella. The exumbrella is divided by eight radial furrows, each of which runs from the apex (tip) to the edges and to the centers of the four sides. The umbrella jelly is a little thinner in the furrows. The furrows are not clearly developed in all species and can also become more indistinct with age. The apex can also be accentuated by an annular furrow. Parts or the entire exumbrella can be covered with warts, which often show a regular pattern. The warts are usually covered with nettle cells. The exumbrella often shows a characteristic color.

The edge of the umbrella is whole, in contrast to the lobed edge of the umbrella in Medusa of Scyphozoa . The (lower) opening of the umbrella can be narrowed by a muscular fold of skin, the velarium. This is formed exclusively from cell layers of the subumbrella, i.e. not directly homologous to the velum of the hydrozoa , which arises from the epidermal layers of exumbrella and subumbrella. The velarium is criss-crossed by velar canals that are more or less branched. The shape and number of the velar canals are often species-specific.

The interior of the medusa ("gastrovascular system") contains the interior of the manubrium, the stomach, the four gastric pockets and a peripheral ring channel from which the channels in the pedals or tentacles, the velarium and the ropes branch off. The flat stomach lies at the upper end or in the apex of the subumbrella. The food can get into the stomach cavity via the wide interior space of the manubrium. From the interradial areas of the stomach wall, tufted, glandular gastric filaments extend into the interior of the stomach. They produce digestive secretions. Four perradial ostia, which can be closed by a flap, connect the inside of the stomach with the flat gastric pockets that lie on the inside of the umbrella sides. These are thus formed by gastrodermal cell layers of exumbrella and subumbrella. The gastric pockets are separated from one another by narrow, interradial, vertical ridges or septa. In the lower part, up to about the level of the Rhopalia, each gastric pocket is divided into two narrow, vertical marginal pockets by protuberances in the wall. However, these eight marginal pockets are connected to one another on the outer edge and thus form an annular channel. The leaf-shaped gonads (sex glands) are located in the upper gastric pockets and attach in pairs to the vertical septa. In Tripedalia cystophora , the eggs are fertilized in the gastric pouches and the larvae developing from the fertilized eggs only leave the gastric space after a few days.

The nervous system consists of a subumbrellar nerve ring, which is located near the edge of the screen. It connects the sense organs (Rhopalia) and the tentacles. The subumbrella also has a diffuse nerve network with numerous ganglion cells.

The tentacles insert at the lower corners of the cube or cuboid, i.e. interradially. However, they do not start directly on the glider , but on intermediate so-called pedals . However, the term is used to refer to two somewhat different structures. The Carybdeida are roughly triangular or oar-like structures, with the oars (blades) arranged perpendicular to the body surface. They are muscular but not contractile. However, they can be bent outwards and inwards and aid in directional swimming and the transport of food to the mouth. The tentacles arise at the ends of the pedalia, one to three in the order Carybdeida. In the order Chirodropida , the tentacles insert around a palm-shaped structure with opposing or offset tentacles and an unpaired outer tentacle that sits at the end of a conical structure and is significantly higher than the other tentacles. It lies on the outside of the other tentacles and covers their beginnings when you look at the pedal from outside. This type is also called "compound pedalium". There are up to around 60 tentacles, around 15 per pedal. The tentacles are hollow, very muscular and highly contractile or highly stretchable. They are usually very long, always significantly longer than the height of the umbrella. They are occupied by thousands of nettle cells (nematocytes or cnidocytes) that contain the nettle capsules (nematocysts or cnidocysts). The nettle capsules are usually not evenly distributed on the tentacles, but rather concentrated in rings. They are arranged perpendicular or slightly obliquely to the surface of the tentacles. They carry different types of stinging cells, which types occur where on the tentacle areas is species-specific and varies accordingly.

Stinging cells

Light microscope image of nettle cells isolated from the tentacles of Chironex fleckeri (400x magnification)

In the box jellyfish, only six different basic types of stomocnid have been observed: three different haplonemes (atric, basic and holotric isorhizas) and three different heteronemes (microbasic mastigophores, heterotrichic microbasic euryteles and stenoteles). However, a further subdivision of these six types has already been achieved in more recent studies, e.g. B. the isorhizas. The nettle cells arise in special formation cells, the nematoblasts or cnidoblasts. They are often only transported to the actual place of their use as so-called wandering cells. They are either stored individually between the epidermal cells or concentrated in batteries of several nettle cells. The discharge occurs through the mechanical stimulation of the cnidocil, a stiff bristle of the nettle cell that has emerged from a flagellum. The nettle cell itself is closed by a cover (operculum). Inside there is a long rolled up hollow tube (thread, or also called nema) with a shaft. The different types of nettle cells are classified according to the different shape and function of shaft and thread. When the cnidocil is mechanically stimulated, the stinging cells explode and detonate the operculum. Due to the overpressure of 140 bar, the shaft and thread turn inside out in about 1/250 of a second. In some types of nettle cells, the shafts are provided with fine stylets, which can easily penetrate the skin, fold outwards as the discharge continues and make the shaft look like a harpoon. The thread is also covered with fine thorns, which initially facilitate penetration, later anchor the thread firmly in the tissue by spreading it. The threads release a poison or a poison cocktail into the victim's skin, which, depending on the type, has cytotoxic , neurotoxic , hemolytic , cardiotoxic , dermatonecrotic , immunogenic and inflammatory effects.

The entire equipment with different types of nettle cells and their arrangement on the screen, manubrium and tentacles is also referred to as a cnidome.

Sense organs

Eye of Carybdea marsupialis . Epi = epidermis; Cor = cornea; Lin = lens (partly crystal clear, partly still with recognizable cells (drawn in red)); Lik = lens capsule; Pri = prism cells; Pyr = pyramidal cells

At the edge of the exumbrella, i.e. on the side surfaces of the cube or cuboid, four club-shaped or piston-shaped, relatively large sense organs, the ropes, sit perradially in pits . Phylogenetically, they arose from transformed tentacles. This can also be observed during ontogenesis . They have a two-layer structure with an outer epidermal layer and an inner gastrodermal layer. The rhopalia are connected to the gastric space by the pedicel canal. A total of 24 eyes sit on the Rhopalia, each with two lens eyes, two slit-shaped pigment eyes and two pit-shaped pigment eyes as well as the statocyst (organ of equilibrium) at the base of the Rhopalium. The pigment eyes are also referred to as secondary eyes. The eyes (and also the statocyst) are virtually embedded in a compact mass of nerve cells. The lens eyes, however, are turned towards the inside of the screen and can only see black and white. In laboratory tests, medusas cleverly avoided dark objects.

The equilibrium organ, the statocyst , is located in the lower part of the rhopalium . It contains a compact statolith made of plaster or basanite . It shows fine growth rings, which are interpreted as daily growth rings.

Nettle poisons and their effects

The nettle poison of the box jellyfish is one of the strongest poisons in the animal kingdom. It is used to catch prey and to defend against enemies. The poison, which is in the nettle capsules of the approximately 1.2 m long tentacles of Chironex fleckeri , causes the death of a child who comes into contact with it within a few minutes. The length of a single tentacle of an adult Chironex fleckeri can reach up to 3 m. An adult specimen has up to 60 tentacles; this results in a theoretical total tentacle length of 180 m. The poison of a single specimen of Chironex fleckeri could theoretically kill over 100 people. According to other sources, six to eight meters of tentacles have enough venom to kill a human. Tourist deaths were reported on the Thai islands of Ko Pha-ngan and Ko Samui in the mid-2010s . Poisoning with nettle poison from another group of box jellyfish ("Irukandji group") triggers the extremely painful Irukandji syndrome , which is very rarely fatal. So far, the toxins and their structures have hardly been elucidated, it is only clear that they consist of proteins.

In the film Sieben Leben plays u. a. a box jellyfish play a role.

Life cycle

Life cycle of a box jellyfish

Reproduction can take place both in the polyp stage and in the medusa stage. In the polyp stage, only asexual reproduction by budding has so far been observed. If the polyp is well fed, several buds can be formed at the same time. They separate individually from the body of the "mother" polyp and crawl away. The polyp takes on a special stage of development, the creeping polyp. The polyp stretches very strongly and moves forward with the contracted tentacles (e.g. Carybdea marsupialis ). The creeping polyp of Tripedalia, on the other hand, stretches out a single tentacle and carries it forward as a search organ. How the polyp moves has not yet been clarified. The creeping phase lasts about two to three days, then the polyp attaches itself to a suitable solid ground and becomes a "normal" polyp, which can reproduce again by budding or can be transformed into a medusa by metamorphosis.

Only sexual reproduction occurs in the medusa stage. The sexes are separate. Outwardly, the polyps do not yet differ in terms of males and females. But even with the medusas they can hardly be distinguished from the outside because of the transparency of all organs. The time it takes to reach sexual maturity is species-specific. In Tripedalia cystophora it is about ten to twelve weeks.

The sex products, eggs and sperm are mostly released into the open water, where fertilization takes place (e.g. with Chironex fleckeri ). In other species, however, internal fertilization was observed. In the males of Tripedalia cystophora , the sperm are grouped into sperm bundles. Several sperm bundles are bundled into round spermatophore spheres in special gastric pockets and surrounded by a membrane. The sperm transfer only takes place after a relatively complex mating game. First males and females swim next to each other. Then the male catches the female's tentacle with a tentacle, after which the partners turn in circles. The male transfers the spermatophores to a tentacle of the female. The partners split up. The female guides the spermatophores to the gastric space with the help of the tentacle. There the shell dissolves and the sperm can fertilize the eggs expelled from the gonads in the gastric pockets. The gonads are almost completely emptied during mating, but can regenerate and further mating can occur (at least under breeding conditions).

The fertilized eggs develop into so-called planula larvae in the gastric pouches in two to three days . These then leave the gastric pockets and continue to develop in the open water (larviparia). After another two to three days, they attach to a suitable solid ground and develop into a polyp. This initially only forms two or three tentacles and thus differs significantly from the primary polyp of the umbrella jellyfish ( Scyphozoa ), in which four tentacles are the first to develop.

Way of life

An alternating filling and emptying of their umbrella cavity enables the animals to move using the recoil principle. They are relatively fast swimmers who swim purposefully, but can also make quick 180 ° turns. Chironex fleckeri moves at up to 414 m / h along the coast. Carybdea marsupialis pulsates 120 to 150 times per minute and covers 3 to 6 m (i.e. 180 to 360 m / h). In terms of swimming and migration behavior, they are more comparable to fish. The “planktonic” way of life of the “jellyfish”, which is often rumored in popular scientific literature, certainly does not apply to the box jellyfish ( Cubozoa ), they belong to the necton .

So far, only a few species have been researched so well that the entire life cycle is known. Usually only the medusa stages are described. The medusas are mostly found in the shallow coastal waters (upper sublittoral ), where they hunt fish and crustaceans. Only one species ( Chirodectes maculatus ) is assumed to be an inhabitant of deeper waters of the shelf . The development of most species is strongly seasonal. In northern Queensland (Australia), Chironex fleckeri appears in October to December and then disappears in April to May. The polyps were meanwhile in mangrove channels of large estuaries z. T. found up to 15 km inland. Under laboratory conditions it was found that they tolerate large fluctuations in salinity relatively well. The polyps can reproduce asexually up to a minimum salt content of 20 per mille, they survive a salt content of about five per mille. The tolerance of higher salt contents was not tested, but no impairments were observed at salt contents around 42 per thousand. So far, little is known how old the medusas can get. For smaller species, the lifespan is likely to be around three to four months. Since some species have an annual cycle and are strongly influenced by the seasons in their habitat, it can be assumed that the animals only live a total of one year. This need not apply to the species that live on the deeper shelf (e.g. Chirodectes maculatus ).

The Medusas of box jellyfish are active predators, which differ from zooplankton , especially copepods (Copepoda), shrimp , amphipods (amphipods), fish larvae and small fish feed.

Geographical distribution

Most species of box jellyfish inhabit the shallow coastal waters of tropical and subtropical waters of the Indic, Pacific and Atlantic. Only a few species also penetrate into temperate latitudes, for example Carybdea marsupialis as far as the Mediterranean and the Portuguese coast. The distribution area of Tamoya haplonema stretches from the Caribbean to Cape Cod (in the southeast of the state of Massachusetts , USA). Carybdea rastonii can be found from the southern tip of Australia to Hokkaidō (Japan). The respective diversity in a marine area is usually not high. There are usually only two to three species in the same range.

Anthracomedusa from the Mazon Creek (Middle Pennsylvania ) deposit of Illinois, USA (FMNH PE 38977, Field Museum of Natural History, Chicago, Illinois, USA)

Fossil box jellyfish

The oldest generally recognized fossil box jellyfish is Anthracomedusa turnbulli from the Pennsylvania of Illinois (USA). Two other fossil box jellyfish, Bipedalia cerinensis and Paracarybdea lithographica , were described in 2006 from the Upper Kimmeridgian (Jura) by Cerin (France). A box jellyfish from the Cambrian is described by Cartwright et al. (2007) in open nomenclature, i.e. without assigning scientific names, from the Middle Cambrian of the Marjum Formation of Utah (USA). With the reinterpretation of the fossil Yunnanoascus haikouensis in 2016, the first species from the Chinese Chengjiang fauna was also described; this would be the oldest box jellyfish.

Systematics

The box jellyfish were long classified as the order Cubomedusae in the umbrella jellyfish (Scyphozoa). Werner (1975) then found serious differences to the umbrella jellyfish and therefore established the class of Cubozoa.

Classic system

The classic hierarchical system has undergone major expansion and modification in recent years through numerous new descriptions and reworkings. It was not until 2010 that Bastian Bentlage et al. modified and introduced some new taxa. At present, two orders are distinguished, Chirodropida and Carybdeida:

Phylogeny

According to the molecular genetic analysis by Collins (2002) and a combined morphological-molecular genetic analysis by Marques & Collins (2004), the box jellyfish (Cubozoa) are the sister taxon of the Staurozoa . Both taxa together are in turn the sister taxon of the umbrella jellyfish (Scyphozoa). Cnidaria cladogram according to Collins (2002)

  Cnidaria  


 Hexacorallia 


   

 Octocorallia 


   


 Scyphozoa 


   

 Staurozoa 


   

 Cubozoa 




   

 Hydrozoa 



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Internal phylogeny according to Bentlage et al. (2010)

  Cubozoa  
  Carybdeida  




 Carybdeidae 


   

 Tripedaliidae 



   

 Tamoyidae 



   

 Carukiidae 



   

 Alatinidae 



  Chirodropida  


 Chirodropidae 


   

 Chiropsellidae



   

 Chiropsalmidae




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Individual evidence

  1. a b I. Straehler-Pohl and G. Jarms: Life cycle of Carybdea marsupialis Linnaeus, 1758 (Cubozoa, Carybdeidae) reveals metamorphosis to be a modified strobilation. Marine Biology, 147: 1271-1277, London 2005 doi : 10.1007 / s00227-005-0031-4
  2. a b Carybdea marsupialis was originally described by Carl von Linné from the Mediterranean. The animals that Bernhard Werner u. a. when Carybdea marsupialis were determined and used for his fundamental work on biology, however, originated in Puerto Rico. The polyp culture, which has been held for 40 years, was also used for other fundamental studies of the biology of the box jellyfish. According to the molecular genetic studies 18S (small subunit) and 28S (large subunit) as well as mitochondrial 16S by Bentlage et al. this culture is very close to Alatina spp. related. There is strong suspicion that this population is not Carybdea marsupialis , but an Alatina species. Alatina spp. occurs in Puerto Rico, while Carybdea marsupialis is otherwise restricted to the Mediterranean.
  3. ^ A b Peter J. Fenner: Dangers in the Ocean: The Traveler and Marine Envenomation. I. Jellyfish. Journal of Travel Medicine, 5 (3): 135-141 1998 doi : 10.1111 / j.1708-8305.1998.tb00487.x
  4. ^ Paul FS Cornelius, Peter J. Fenner and Russell Hore: Chiropsalmus maculatus sp. nov., a cubomedusa from the Great Barrier Reef. Memoirs of the Queensland Museum, 51 (2): 399-405, Brisbane, 2005 ISSN  0079-8835
  5. A. Garm, MM Coates, R. Gad, J. Seymour and D.-E. Nilsson: The lens eyes of the box jelly fish Tripedalia cystophora and Chiropsalmus sp. are slow and color-blind. Journal of Comparative Physiology A, 193: 547-557, 2007 doi : 10.1007 / s00359-007-0211-4
  6. Henry Tiemann, Ilka Sötje, Alexander Becker, Gerhard Jarms and Matthias Epple: Calcium sulfate hemihydrate (bassanite) statoliths in the cubozoan Carybdea sp. Zoologischer Anzeiger - Journal of Comparative Zoology, 245 (1): 13-17, Munich 2006 doi : 10.1016 / j.jcz.2006.03.001
  7. Shunshiro Ueno, Chifumi Imai and Atsushi Mitsutani: Fine growth rings found in statolith of a cubomedusa Carybdea rastoni. Journal of Plankton Research, 17 (6)): 1381-1384, Oxford 1995 ISSN  0142-7873 abstract
  8. after Philipp Alderslade
  9. Thailand: German tourist dies after a box jellyfish sting. SpiegelOnline, October 7, 2015, accessed October 9, 2015 .
  10. ^ MR Gordon and JE Seymour: Quantifying movement of the tropical Australian cubozoan Chironex fleckeri using acoustic telemetry. Hydrobiologia, 616: 87-97, Dordrecht 2009 doi : 10.1007 / s10750-008-9594-7
  11. ^ RF Hartwick: Distributional ecology and behavior of the early life stages of the box-jellyfish Chironex fleckeri . Hydrobiologia, 216/217: 181-188, Dordrecht 1991 ISSN  0018-8158 PDF
  12. Christian Gaillard, Jaqueline Goy, Paul Bernier, Jean Paul Bourseau, Jean Claude Gall, Georges Barale, Eric Buffetaut and Sylvie Wenz: New jellyfish taxa from the Upper Jurassic Lithographic Limestones of Cerin (France): Taphonomy and Ecology. Palaeontology, 49 (6): 1287-1302, London 2006
  13. ^ Paulyn Cartwright, Susan L. Halgedahl et al. a .: Exceptionally Preserved Jellyfishes from the Middle Cambrian. In: PLoS ONE. 2, 2007, p. E1121, doi : 10.1371 / journal.pone.0001121 .
  14. Jian Han, Shixue Hu, Paulyn Cartwright, Fangchen Zhao, Qiang Ou, Shin Kubota, Xing Wang, Xiaoguang Yang (2016): The earliest pelagic jellyfish with rhopalia from Cambrian Chengjiang deposit. Palaeogeography, Palaeoclimatology, Palaeoecology 449: 166-173. doi : 10.1016 / j.palaeo.2016.02.025
  15. Sho Toshino, Hiroshi Miyake, Haruka Shibata: Meteorona kishinouyei, a new family, genus and species (Cnidaria, Cubozoa, Chirodropida) from Japanese Waters. Zookeys, 503: 1-21, 2015 doi : 10.3897 / zookeys.503.9047
  16. Bastian Bentlage: Description of the chirodropid box jellyfish Chiropsella rudloei sp. nov. (Cnidaria: Cubozoa) from Madagascar. Marine Biodiversity Records, 2013 PDF
  17. Lisa-Ann Gershwin, Merrick Ekins: A new pygmy species of box jellyfish (Cubozoa: Chirodropida) from sub-tropical Australia. Marine Biodiversity Records, 8, 2015, e111 (8 pages) doi : 10.1017 / S175526721500086X
  18. a b c Lisa-Ann Gershwin: Two new species of box jellies (Cnidaria: Cubozoa: Carybdeida) from the central coast of Western Australia, both presumed to cause Irukandji syndrome. Records of the Western Australian Museum 29 (1): 10-19, 2014 PDF
  19. Bastian Bentlage, Cheryl Lewis: An illustrated key and synopsis of the families and genera of carybdeid box jellyfishes (Cnidaria: Cubozoa: Carybdeida), with emphasis on the “Irukandji family” (Carukiidae). Journal of Natural History, 46 (41-42): 2595-2620, 2012, doi : 10.1080 / 00222933.2012.717645
  20. Allen G. Collins, Bastian Bentlage, William Gillan, Tara H. Lynn, André C. Morandini and Antonio C. Marques: Naming the Bonaire banded box jelly, Tamoya ohboya, n. Sp. (Cnidaria: Cubozoa: Carybdeida: Tamoyidae). Zootaxa, 2753: 53-68, 2011 PDF

literature

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