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(Platyhelminthes)

Flatworms

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Platyhelminthes is a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrates commonly called flatworms or flat worms. Being acoelomates (having no body cavity), and having no specialised circulatory and respiratory organs, they are restricted to having flattened shapes that allow oxygen and nutrients to pass through their bodies by diffusion. The digestive cavity has only one opening for both ingestion (intake of nutrients) and egestion (removal of undigested wastes); as a result, the food can not be processed continuously.

Free-living flatworms are mostly predators, and live in water or in shaded, humid terrestrial environments, such as leaf litter. Cestodes (tapeworms) and trematodes (flukes) have complex life-cycles, with mature stages that live as parasites in the digestive systems of fish or land vertebrates, and intermediate stages that infest secondary hosts. The eggs of trematodes are excreted from their main hosts, whereas adult cestodes generate vast numbers of hermaphroditic, segment-like proglottids that detach when mature, are excreted, and then release eggs. Unlike the other parasitic groups, the monogeneans are external parasites infesting aquatic animals, and their larvae metamorphose into the adult form after attaching to a suitable host.

The lack of circulatory and respiratory organs limits platyhelminths to sizes and shapes that enable oxygen to reach and carbon dioxide to leave all parts of their bodies by simple diffusion. Hence, many are microscopic, and the large species have flat ribbon-like or leaf-like shapes. Because there is no circulatory system which can transport nutrients around, the guts of large species have many branches, allowing the nutrients to diffuse to all parts of the body. Respiration through the whole surface of the body makes them vulnerable to fluid loss, and restricts them to environments where dehydration is unlikely: sea and freshwater, moist terrestrial environments such as leaf litter or between grains of soil, and as parasites within other animals.

The space between the skin and gut is filled with mesenchyme, also known as parenchyma, a connective tissue made of cells and reinforced by collagen fibers that act as a type of skeleton, providing attachment points for muscles. The mesenchyme contains all the internal organs and allows the passage of oxygen, nutrients and waste products. It consists of two main types of cell: fixed cells, some of which have fluid-filled vacuoles; and stem cells, which can transform into any other type of cell, and are used in regenerating tissues after injury or asexual reproduction.

Most platyhelminths have no anus and regurgitate undigested material through the mouth. The genus Paracatenula, whose members include tiny flatworms living in symbiosis with bacteria, is even missing a mouth and a gut. However, some long species have an anus and some with complex, branched guts have more than one anus, since excretion only through the mouth would be difficult for them. The gut is lined with a single layer of endodermal cells that absorb and digest food. Some species break up and soften food first by secreting enzymes in the gut or pharynx (throat).

All animals need to keep the concentration of dissolved substances in their body fluids at a fairly constant level. Internal parasites and free-living marine animals live in environments with high concentrations of dissolved material, and generally let their tissues have the same level of concentration as the environment, while freshwater animals need to prevent their body fluids from becoming too dilute. Despite this difference in environments, most platyhelminths use the same system to control the concentration of their body fluids. Flame cells, so called because the beating of their flagella looks like a flickering candle flame, extract from the mesenchyme water that contains wastes and some reusable material, and drive it into networks of tube cells which are lined with flagella and microvilli. The tube cells’ flagella drive the water towards exits called nephridiopores, while their microvilli reabsorb reusable materials and as much water as is needed to keep the body fluids at the right concentration. These combinations of flame cells and tube cells are called protonephridia.

In all platyhelminths, the nervous system is concentrated at the head end. Other platyhelminths have rings of ganglia in the head and main nerve trunks running along their bodies.

Source: Wikipedia

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(Xenacoelomorpha)

Xenacoelomorphs

Ксенакоеломорфи

Xenacoelomorpha is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies (physical appearances shared by the animals in the clade), which was then confirmed by phylogenomic analyses of molecular data (similarities in the DNA of the animals within the clade).

The phylum consists of small, flat and worm-like creatures found in marine and sometimes brackish water environments, on the sediments. There are species that are variously free-living, parasitic, and symbiotic. They can be found at depths of almost 4 km and near hydrothermal vents.

The phylum is hermaphroditic (all individuals have both male and female sex organs) and reproduces sexually with direct development, meaning they skip a potentially vulnerable larval stage. Xenoturbella have external fertilization, and Acoelomorpha has internal fertilization. All xenacoelomorphs are bilateral, meaning they have a central front-to-back body axis with mirror image right and left sides. They are triploblasts (meaning they have the three germ layers: ectoderm, endoderm, and mesoderm). Their body plan is acoelomate – they lack a coelom – do not have a true body cavity. Also an excretory system is absent, yet all genes related to the excretory system are present except for Osr, which is essential for the development for such a system. In acoelomorphs, which has gone through rapid evolutionary rates and chromosomic rearrangements, about 60% of the genes shared between protostomes and deuterostomes are missing. How many of these genes which are present or absent in Xenoturbella will require a whole genome sequencing.

While other animals that are diploblastic (only have two germ layers: ectoderm and endoderm) also lack a coelom, those technically do not have an acoelomate body plan because they lack the mesoderm germ layer. In acoels, the mouth opens directly into a large endodermal syncytium, while in nemertodermatids and xenoturbellids there is a sack-like gut lined by unciliated cells.

A defining feature is a digestive system lacking nerve cells. Because an enteric nervous system, also called the stomatogastric nervous system, is also found in many cnidarians, its absence is most likely a derived trait.

Their nervous systems are basiepidermal – located right under the epidermis – and they have no brain. The xenoturbellids’ nervous system consists of a simple nerve net, with no special concentration of neurons. In acoelomorphs the nervous system is arranged in a series of longitudinal bundles, united in the anterior region by a ring comissure of variable complexity.

The sensory organs include a statocyst (for balance). Some groups have two unicellular ocelli (simple eyes).

The epidermis of all species within the phylum is ciliated. The cilia are composed of a set of nine pairs of peripheral microtubules and one or two central microtubules (patterns 9+1 and 9+2, respectively). The pairs 4–7 terminate before the tip, creating a structure called a “shelf”.

Source: Wikipedia

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(Acoela)

Acoels

Ацели

Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order of turbellarian flatworms. About 400 species are known, but probably many more not yet described.

Acoels are very small flattened worms, usually under 2 mm in length, but some larger species, such as Symsagittifera roscoffensis, may reach up to 15 mm. They are bilaterally symmetric and microscopic.

They are found worldwide in marine and brackish waters, usually having a benthic lifestyle, although some species are epibionts. Two species, Limonoposthia polonica and Oligochoerus limnophilus, live in freshwater.

Species in the family Convolutidae often form endosymbiotic relationship with microalgae. In one of the genera, Waminoa, the algae are transmitted vertically from parents to offspring. In addition to Convolutidae, there appears to be a potential new and yet unnamed family of acoels that also live in relationships with microalgal endosymbionts.

Members of the class Acoela lack a conventional gut, so that the mouth opens directly into the mesenchyme, i.e., the layer of tissue that fills the body. Digestion is accomplished by means of a syncytium that forms a vacuole around ingested food. There are no epithelial cells lining the digestive vacuole, but in the families Diopisthoporidae, Hallangiidae, Hofsteniidae, and Solenofilomorphidae, and the genera Oligochoerus (Convolutidae) and Proporus (Proporidae), there is a short pharynx leading from the mouth to the vacuole. All other bilateral animals (apart from tapeworms) have a gut lined with epithelial cells. As a result, the acoels appear to be solid-bodied.

As the basal lineage of bilateral animals, the Acoela provide interesting insights into early animal evolution and development. The most thoroughly studied animal in this group is the species Isodiametra pulchra. Acoela used to be classified in the phylum Platyhelminthes. However, Acoela was separated from this phylum after molecular analyses showed that it had diverged before the three main bilaterian clades had formed, making flatworms as traditionally understood an evolutionary grade from which higher animals had evolved.

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(Ctenophora)

Comb Jellies

Реброплави

Ctenophora comprise 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 are the largest animals to swim with the help of cilia.

Depending on the species, adult ctenophores range from a few millimeters to 1.5 m (5 ft) in size. 186 living species are recognised.

Their bodies consist of a mass of jelly, with a layer two cells thick on the outside, and another lining the internal cavity. The phylum has a wide range of body forms, including the egg-shaped cydippids with a pair of retractable tentacles that capture prey, the flat, generally combless platyctenids, and the large-mouthed beroids, which prey on other ctenophores.

Almost all ctenophores function as predators, taking prey ranging from microscopic larvae and rotifers to the adults of small crustaceans; the exceptions are juveniles of two species, which live as parasites on the salps on which adults of their species feed.

Feeding, Excretion and Respiration

When prey is swallowed, it is liquefied in the pharynx by enzymes and by muscular contractions of the pharynx. The resulting slurry is wafted through the canal system by the beating of the cilia, and digested by the nutritive cells. The ciliary rosettes in the canals may help to transport nutrients to muscles in the mesoglea. The anal pores may eject unwanted small particles, but most unwanted matter is regurgitated via the mouth.

Little is known about how ctenophores get rid of waste products produced by the cells. The ciliary rosettes in the gastrodermis may help to remove wastes from the mesoglea, and may also help to adjust the animal’s buoyancy by pumping water into or out of the mesoglea.

Locomotion

The outer surface bears usually eight comb rows, called swimming-plates, which are used for swimming. The rows are oriented to run from near the mouth (the “oral pole”) to the opposite end (the “aboral pole”), and are spaced more or less evenly around the body, although spacing patterns vary by species and in most species the comb rows extend only part of the distance from the aboral pole towards the mouth. The “combs” (also called “ctenes” or “comb plates”) run across each row, and each consists of thousands of unusually long cilia, up to 2 mm. Unlike conventional cilia and flagella, which has a filament structure arranged in a 9 + 2 pattern, these cilia are arranged in a 9 + 3 pattern, where the extra compact filament is suspected to have a supporting function. These normally beat so that the propulsion stroke is away from the mouth, although they can also reverse direction. Hence ctenophores usually swim in the direction in which the mouth is eating, unlike jellyfish. When trying to escape predators, one species can accelerate to six times its normal speed; some other species reverse direction as part of their escape behavior, by reversing the power stroke of the comb plate cilia.

It is uncertain how ctenophores control their buoyancy, but experiments have shown that some species rely on osmotic pressure to adapt to the water of different densities. Their body fluids are normally as concentrated as seawater. If they enter less dense brackish water, the ciliary rosettes in the body cavity may pump this into the mesoglea to increase its bulk and decrease its density, to avoid sinking. Conversely, if they move from brackish to full-strength seawater, the rosettes may pump water out of the mesoglea to reduce its volume and increase its density.

Reproduction and Development

Adults of most species can regenerate tissues that are damaged or removed, although only platyctenids reproduce by cloning, splitting off from the edges of their flat bodies fragments that develop into new individuals. Lab research on Mnemiopsis leidyi also show that when two individuals have parts of their bodies removed, they are able to fuse together, including their nervous and digestive systems, even when the two individuals are genetically different; a phenomenon that has so far only been found in comb jellies.

The last common ancestor (LCA) of the ctenophores was hermaphroditic. Some are simultaneous hermaphrodites, which can produce both eggs and sperm at the same time, while others are sequential hermaphrodites, in which the eggs and sperm mature at different times. There is no metamorphosis. At least three species are known to have evolved separate sexes (dioecy); Ocyropsis crystallina and Ocyropsis maculata in the genus Ocyropsis and Bathocyroe fosteri in the genus Bathocyroe. The gonads are located in the parts of the internal canal network under the comb rows, and eggs and sperm are released via pores in the epidermis. Fertilization is generally external, but platyctenids use internal fertilization and keep the eggs in brood chambers until they hatch. Self-fertilization has occasionally been seen in species of the genus Mnemiopsis, and it is thought that most of the hermaphroditic species are self-fertile.

Development of the fertilized eggs is direct; there is no distinctive larval form. Juveniles of all groups are generally planktonic, and most species resemble miniature adult cydippids, gradually developing their adult body forms as they grow. In the genus Beroe, however, the juveniles have large mouths and, like the adults, lack both tentacles and tentacle sheaths. In some groups, such as the flat, bottom-dwelling platyctenids, the juveniles behave more like true larvae. They live among the plankton and thus occupy a different ecological niche from their parents, only attaining the adult form by a more radical ontogeny after dropping to the sea-floor.

At least in some species, juvenile ctenophores appear capable of producing small quantities of eggs and sperm while they are well below adult size, and adults produce eggs and sperm for as long as they have sufficient food. If they run short of food, they first stop producing eggs and sperm, and then shrink in size. When the food supply improves, they grow back to normal size and then resume reproduction. These features make ctenophores capable of increasing their populations very quickly. Members of the Lobata and Cydippida also have a reproduction form called dissogeny; two sexually mature stages, first as larva and later as juveniles and adults. During their time as larva they are capable of releasing gametes periodically. After their first reproductive period is over they will not produce more gametes again until later. A population of Mertensia ovum in the central Baltic Sea have become paedogenetic, and consist solely of sexually mature larvae less than 1.6 mm.

In Mnemiopsis leidyi, nitric oxide (NO) signaling is present both in adult tissues and differentially expressed in later embryonic stages suggesting the involvement of NO in developmental mechanisms. The mature form of the same species is also able to revert back to the cydippid stage when triggered by environmental stressors.

Ecology

Ctenophores are found in most marine environments: from polar waters at −2 °C to the tropics at 30 °C; near coasts and in mid-ocean; from the surface waters to the ocean depths at more than 7000 meters. The best-understood are the genera Pleurobrachia, Beroe and Mnemiopsis, as these planktonic coastal forms are among the most likely to be collected near shore. No ctenophores have been found in fresh water.

Ctenophores may be abundant during the summer months in some coastal locations, but in other places, they are uncommon and difficult to find.

In bays where they occur in very high numbers, predation by ctenophores may control the populations of small zooplanktonic organisms such as copepods, which might otherwise wipe out the phytoplankton (planktonic plants), which are a vital part of marine food chains.

Prey and Predators

Almost all ctenophores are predators – there are no vegetarians and only one genus that is partly parasitic. If food is plentiful, they can eat 10 times their own weight per day. While Beroe preys mainly on other ctenophores, other surface-water species prey on zooplankton (planktonic animals) ranging in size from the microscopic, including mollusc and fish larvae, to small adult crustaceans such as copepods, amphipods, and even krill. Members of the genus Haeckelia prey on jellyfish and incorporate their prey’s nematocysts (stinging cells) into their own tentacles instead of colloblasts. Ctenophores have been compared to spiders in their wide range of techniques for capturing prey – some hang motionless in the water using their tentacles as “webs”, some are ambush predators like Salticid jumping spiders, and some dangle a sticky droplet at the end of a fine thread, as bolas spiders do. This variety explains the wide range of body forms in a phylum with rather few species. The two-tentacled “cydippid” Lampea feeds exclusively on salps, close relatives of sea-squirts that form large chain-like floating colonies, and juveniles of Lampea attach themselves like parasites to salps that are too large for them to swallow. Members of the cydippid genus Pleurobrachia and the lobate Bolinopsis often reach high population densities at the same place and time because they specialize in different types of prey: Pleurobrachia’s long tentacles mainly capture relatively strong swimmers such as adult copepods, while Bolinopsis generally feeds on smaller, weaker swimmers such as rotifers and mollusc and crustacean larvae.

Ctenophores used to be regarded as “dead ends” in marine food chains because it was thought their low ratio of organic matter to salt and water made them a poor diet for other animals. It is also often difficult to identify the remains of ctenophores in the guts of possible predators, although the combs sometimes remain intact long enough to provide a clue. Detailed investigation of chum salmon, Oncorhynchus keta, showed that these fish digest ctenophores 20 times as fast as an equal weight of shrimps, and that ctenophores can provide a good diet if there are enough of them around. Beroids prey mainly on other ctenophores. Some jellyfish and turtles eat large quantities of ctenophores, and jellyfish may temporarily wipe out ctenophore populations. Since ctenophores and jellyfish often have large seasonal variations in population, most fish that prey on them are generalists and may have a greater effect on populations than the specialist jelly-eaters. This is underlined by an observation of herbivorous fishes deliberately feeding on gelatinous zooplankton during blooms in the Red Sea. The larvae of some sea anemones are parasites on ctenophores, as are the larvae of some flatworms that parasitize fish when they reach adulthood.

Source: Wikipedia

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(Cnidaria)

Cnidarians

Кнідарії

Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in freshwater and marine environments (predominantly the latter), including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are an uncentralized 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 few animals that can reproduce both sexually and asexually.

Cnidarians mostly have two basic body forms: swimming medusae and sessile polyps, both of which are radially symmetrical with mouths surrounded by tentacles that bear cnidocytes, which are specialized stinging cells used to capture prey. Both forms have a single orifice and body cavity that are used for digestion and respiration. Many cnidarian species produce colonies that are single organisms composed of medusa-like or polyp-like zooids, or both (hence they are trimorphic). Cnidarians’ activities are coordinated by a decentralized nerve net and simple receptors. Cnidarians also have rhopalia, which are involved in gravity sensing and sometimes chemoreception. Several free-swimming species of Cubozoa and Scyphozoa possess balance-sensing statocysts, and some have simple eyes. Not all cnidarians reproduce sexually, but many species have complex life cycles of asexual polyp stages and sexual medusae stages. Some, however, omit either the polyp or the medusa stage, and the parasitic classes evolved to have neither form.

Cnidarians were formerly grouped with ctenophores, also known as comb jellies, in the phylum Coelenterata, but increasing awareness of their differences caused them to be placed in separate phyla. Cnidarians are classified into four main groups: the almost wholly sessile Anthozoa (sea anemones, corals, sea pens); swimming Scyphozoa (jellyfish); Cubozoa (box jellies); and Hydrozoa (a diverse group that includes all the freshwater cnidarians as well as many marine forms, and which has both sessile members, such as Hydra, and colonial swimmers (such as the Portuguese man o’ war)). Staurozoa have recently been recognised as a class in their own right rather than a sub-group of Scyphozoa, and the highly derived parasitic Myxozoa and Polypodiozoa were firmly recognized as cnidarians only in 2007.

Most cnidarians prey on organisms ranging in size from plankton to animals several times larger than themselves, but many obtain much of their nutrition from symbiotic dinoflagellates, and a few are parasites. Many are preyed on by other animals including starfish, sea slugs, fish, turtles, and even other cnidarians. Many scleractinian corals—which form the structural foundation for coral reefs—possess polyps that are filled with symbiotic photo-synthetic zooxanthellae. While reef-forming corals are almost entirely restricted to warm and shallow marine waters, other cnidarians can be found at great depths, in polar regions, and in freshwater.

Description

Most adult cnidarians appear as either free-swimming medusae or sessile polyps, and many hydrozoans species are known to alternate between the two forms.

Both are radially symmetrical, like a wheel and a tube respectively. Since these animals have no heads, their ends are described as “oral” (nearest the mouth) and “aboral” (furthest from the mouth).

Most have fringes of tentacles equipped with cnidocytes around their edges, and medusae generally have an inner ring of tentacles around the mouth. Some hydroids may consist of colonies of zooids that serve different purposes, such as defence, reproduction and catching prey. The mesoglea of polyps is usually thin and often soft, but that of medusae is usually thick and springy, so that it returns to its original shape after muscles around the edge have contracted to squeeze water out, enabling medusae to swim by a sort of jet propulsion.

In medusae, the only supporting structure is the mesoglea. Hydra and most sea anemones close their mouths when they are not feeding, and the water in the digestive cavity then acts as a hydrostatic skeleton, rather like a water-filled balloon. Other polyps such as Tubularia use columns of water-filled cells for support. Sea pens stiffen the mesoglea with calcium carbonate spicules and tough fibrous proteins, rather like sponges.

In some colonial polyps, a chitinous epidermis gives support and some protection to the connecting sections and to the lower parts of individual polyps. A few polyps collect materials such as sand grains and shell fragments, which they attach to their outsides. Some colonial sea anemones stiffen the mesoglea with sediment particles.

A mineralized exoskeleton made of calcium carbonate is found in subphylum Anthozoa in the order Scleractinia (stony corals; class Hexacorallia) and the class Octocorallia, and in subphylum Medusozoa in three hydrozoan families in order Anthoathecata; Milleporidae, Stylasteridae and Hydractiniidae (the latter with a mix of calcified and uncalcified species).

Locomotion

Medusae swim by a form of jet propulsion: muscles, especially inside the rim of the bell, squeeze water out of the cavity inside the bell, and the springiness of the mesoglea powers the recovery stroke. Since the tissue layers are very thin, they provide too little power to swim against currents and just enough to control movement within currents.

Hydras and some sea anemones can move slowly over rocks and sea or stream beds by various means: creeping like snails, crawling like inchworms, or by somersaulting. A few can swim clumsily by waggling their bases.

Feeding and excretion

Cnidarians feed in several ways: predation, absorbing dissolved organic chemicals, filtering food particles out of the water, obtaining nutrients from symbiotic algae within their cells, and parasitism. Most obtain the majority of their food from predation but some, including the corals Hetroxenia and Leptogorgia, depend almost completely on their endosymbionts and on absorbing dissolved nutrients. Cnidaria give their symbiotic algae carbon dioxide, some nutrients, and protection against predators.

Predatory species use their cnidocytes to poison or entangle prey, and those with venomous nematocysts may start digestion by injecting digestive enzymes. The “smell” of fluids from wounded prey makes the tentacles fold inwards and wipe the prey off into the mouth. In medusae, the tentacles around the edge of the bell are often short and most of the prey capture is done by “oral arms”, which are extensions of the edge of the mouth and are often frilled and sometimes branched to increase their surface area. These “oral arms” aid in cnidarians’ ability to move prey towards their mouth once it has been poisoned and entangled. Medusae often trap prey or suspended food particles by swimming upwards, spreading their tentacles and oral arms and then sinking. In species for which suspended food particles are important, the tentacles and oral arms often have rows of cilia whose beating creates currents that flow towards the mouth, and some produce nets of mucus to trap particles. Their digestion is both intra and extracellular.

Once the food is in the digestive cavity, gland cells in the gastroderm release enzymes that reduce the prey to slurry, usually within a few hours. This circulates through the digestive cavity and, in colonial cnidarians, through the connecting tunnels, so that gastroderm cells can absorb the nutrients. Absorption may take a few hours, and digestion within the cells may take a few days. The circulation of nutrients is driven by water currents produced by cilia in the gastroderm or by muscular movements or both, so that nutrients reach all parts of the digestive cavity. Nutrients reach the outer cell layer by diffusion or, for animals or zooids such as medusae which have thick mesogleas, are transported by mobile cells in the mesoglea.

Indigestible remains of prey are expelled through the mouth. The main waste product of cells’ internal processes is ammonia, which is removed by the external and internal water currents.

Regeneration

All cnidarians can regenerate, allowing them to recover from injury and to reproduce asexually. Medusae have limited ability to regenerate, but polyps can do so from small pieces or even collections of separated cells. This enables corals to recover even after apparently being destroyed by predators.

Reproduction

Cnidarian sexual reproduction often involves a complex life cycle with both polyp and medusa stages. For example, in Scyphozoa (jellyfish) and Cubozoa (box jellies), a larva swims until it finds a good site, and then becomes a polyp. This grows normally but then absorbs its tentacles and splits horizontally into a series of disks that become juvenile medusae, a process called strobilation. The juveniles swim off and slowly grow to maturity, while the polyp re-grows and may continue strobilating periodically. The adult medusae have gonads in the gastroderm, and these release ova and sperm into the water in the breeding season.

This phenomenon of succession of differently organized generations (one asexually reproducing, sessile polyp, followed by a free-swimming medusa or a sessile polyp that reproduces sexually) is sometimes called “alternation of asexual and sexual phases” or “metagenesis”, but should not be confused with the alternation of generations as found in plants.

Shortened forms of this life cycle are common, for example some oceanic scyphozoans omit the polyp stage completely, and cubozoan polyps produce only one medusa. Hydrozoa have a variety of life cycles. Some have no polyp stages and some (e.g. hydra) have no medusae. In some species, the medusae remain attached to the polyp and are responsible for sexual reproduction; in extreme cases these reproductive zooids may not look much like medusae. Meanwhile, life cycle reversal, in which polyps are formed directly from medusae without the involvement of sexual reproduction process, was observed in both Hydrozoa (Turritopsis dohrnii and Laodicea undulata) and Scyphozoa (Aurelia). Anthozoa have no medusa stage at all and the polyps are responsible for sexual reproduction.

Spawning is generally driven by environmental factors such as changes in the water temperature, and their release is triggered by lighting conditions such as sunrise, sunset or the phase of the moon. Many species of Cnidaria may spawn simultaneously in the same location, so that there are too many ova and sperm for predators to eat more than a tiny percentage — one famous example is the Great Barrier Reef, where at least 110 corals and a few non-cnidarian invertebrates produce enough gametes to turn the water cloudy. These mass spawnings may produce hybrids, some of which can settle and form polyps, but it is not known how long these can survive. In some species the ova release chemicals that attract sperm of the same species.

The fertilized eggs develop into larvae by dividing until there are enough cells to form a hollow sphere (blastula) and then a depression forms at one end (gastrulation) and eventually becomes the digestive cavity. However, in cnidarians the depression forms at the end further from the yolk (at the animal pole), while in bilaterians it forms at the other end (vegetal pole). The larvae, called planulae, swim or crawl by means of cilia. They are cigar-shaped but slightly broader at the “front” end, which is the aboral, vegetal-pole end and eventually attaches to a substrate if the species has a polyp stage.

Anthozoan larvae either have large yolks or are capable of feeding on plankton, and some already have endosymbiotic algae that help to feed them. Since the parents are immobile, these feeding capabilities extend the larvae’s range and avoid overcrowding of sites. Scyphozoan and hydrozoan larvae have little yolk and most lack endosymbiotic algae, and therefore have to settle quickly and metamorphose into polyps. Instead, these species rely on their medusae to extend their ranges.

All known cnidarians can reproduce asexually by various means, in addition to regenerating after being fragmented. Hydrozoan polyps only bud, while the medusae of some hydrozoans can divide down the middle. Scyphozoan polyps can both bud and split down the middle. In addition to both of these methods, Anthozoa can split horizontally just above the base. Asexual reproduction makes the daughter cnidarian a clone of the adult. The ability of cnidarians to asexually reproduce ensures a greater number of mature medusa that can mature to reproduce sexually.

Ecology

Many cnidarians are limited to shallow waters because they depend on endosymbiotic algae for much of their nutrients. The life cycles of most have polyp stages, which are limited to locations that offer stable substrates. Nevertheless, major cnidarian groups contain species that have escaped these limitations. Hydrozoans have a worldwide range: some, such as Hydra, live in freshwater; Obelia appears in the coastal waters of all the oceans; and Liriope can form large shoals near the surface in mid-ocean. Among anthozoans, a few scleractinian corals, sea pens and sea fans live in deep, cold waters, and some sea anemones inhabit polar seabeds while others live near hydrothermal vents over 10 km below sea-level. Reef-building corals are limited to tropical seas between 30°N and 30°S with a maximum depth of 46 m, temperatures between 20 and 28 °C, high salinity, and low carbon dioxide levels. Stauromedusae, although usually classified as jellyfish, are stalked, sessile animals that live in cool to Arctic waters. Cnidarians range in size from a mere handful of cells for the parasitic myxozoans through Hydra’s length of 5–20 mm, to the lion’s mane jellyfish, which may exceed 2 m in diameter and 75 m in length.

Prey of cnidarians ranges from plankton to animals several times larger than themselves. Some cnidarians are parasites, mainly on jellyfish but a few are major pests of fish. Others obtain most of their nourishment from endosymbiotic algae or dissolved nutrients. Predators of cnidarians include: sea slugs, flatworms and comb jellies, which can incorporate nematocysts into their own bodies for self-defense (nematocysts used by cnidarian predators are referred to as kleptocnidae); starfish, notably the crown of thorns starfish, which can devastate corals; butterfly fish and parrot fish, which eat corals; and marine turtles, which eat jellyfish. Some sea anemones and jellyfish have a symbiotic relationship with some fish; for example clownfish live among the tentacles of sea anemones, and each partner protects the other against predators.

Coral reefs form some of the world’s most productive ecosystems. Common coral reef cnidarians include both anthozoans (hard corals, octocorals, anemones) and hydrozoans (fire corals, lace corals). The endosymbiotic algae of many cnidarian species are very effective primary producers, in other words converters of inorganic chemicals into organic ones that other organisms can use, and their coral hosts use these organic chemicals very efficiently. In addition, reefs provide complex and varied habitats that support a wide range of other organisms. Fringing reefs just below low-tide level also have a mutually beneficial relationship with mangrove forests at high-tide level and seagrass meadows in between: the reefs protect the mangroves and seagrass from strong currents and waves that would damage them or erode the sediments in which they are rooted, while the mangroves and seagrass protect the coral from large influxes of silt, fresh water and pollutants. This additional level of variety in the environment is beneficial to many types of coral reef animals, which for example may feed in the sea grass and use the reefs for protection or breeding.

Source: Wikipedia

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(Porifera)

Sponges

Губки

Sponges or sea sponges are primarily marine invertebrates of the metazoan phylum Porifera, a basal animal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and are one of the most ancient members of macrobenthos, with many historical species being important reef-building organisms.

Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues and organs. Some of them are radially symmetrical, but most are asymmetrical. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where the water deposits nutrients and then leaves through a hole called the osculum. The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have been used as morphological evidence to suggest sponges are the sister group to the rest of animals. A great majority are marine (salt-water) species, ranging in habitat from tidal zones to depths exceeding 8,800 m, though there are freshwater species. All adult sponges are sessile, meaning that they attach to an underwater surface and remain fixed in place (i.e., do not travel). While in their larval stage of life, they are motile.

Many sponges have internal skeletons of spicules (skeletal-like fragments of calcium carbonate or silicon dioxide), and/or spongin (a modified type of collagen protein). An internal gelatinous matrix called mesohyl functions as an endoskeleton, and it is the only skeleton in soft sponges that encrust such hard surfaces as rocks. More commonly, the mesohyl is stiffened by mineral spicules, by spongin fibers, or both. 90% of all known sponge species that have the widest range of habitats including all freshwater ones are demosponges that use spongin; many species have silica spicules, whereas some species have calcium carbonate exoskeletons. Calcareous sponges have calcium carbonate spicules and, in some species, calcium carbonate exoskeletons, are restricted to relatively shallow marine waters where production of calcium carbonate is easiest.  The fragile glass sponges, with “scaffolding” of silica spicules, are restricted to polar regions and the ocean depths where predators are rare. Fossils of all of these types have been found in rocks dated from 580 million years ago. In addition Archaeocyathids, whose fossils are common in rocks from 530 to 490 million years ago, are now regarded as a type of sponge.

Although most of the approximately 5,000–10,000 known species of sponges feed on bacteria and other microscopic food in the water, some host photosynthesizing microorganisms as endosymbionts, and these alliances often produce more food and oxygen than they consume. A few species of sponges that live in food-poor environments have evolved as carnivores that prey mainly on small crustaceans.

Most sponges reproduce sexually, but they can also reproduce asexually. Sexually reproducing species release sperm cells into the water to fertilize ova released or retained by its mate or “mother”; the fertilized eggs develop into larvae which swim off in search of places to settle.  Sponges are known for regenerating from fragments that are broken off, although this only works if the fragments include the right types of cells. Some species reproduce by budding. When environmental conditions become less hospitable to the sponges, for example as temperatures drop, many freshwater species and a few marine ones produce gemmules, “survival pods” of unspecialized cells that remain dormant until conditions improve; they then either form completely new sponges or recolonize the skeletons of their parents.

Habitats

Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the polar regions to the tropics. Most live in quiet, clear waters, because sediment stirred up by waves or currents would block their pores, making it difficult for them to feed and breathe. The greatest numbers of sponges are usually found on firm surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like base.

Sponges are more abundant but less diverse in temperate waters than in tropical waters, possibly because organisms that prey on sponges are more abundant in tropical waters. Glass sponges are the most common in polar waters and in the depths of temperate and tropical seas, as their very porous construction enables them to extract food from these resource-poor waters with the minimum of effort. Demosponges and calcareous sponges are abundant and diverse in shallower non-polar waters.

Life Cycle

Sponges in temperate regions live for at most a few years, but some tropical species and perhaps some deep-ocean ones may live for 200 years or more. Some calcified demosponges grow by only 0.2 mm per year and, if that rate is constant, specimens 1 m wide must be about 5,000 years old. Some sponges start sexual reproduction when only a few weeks old, while others wait until they are several years old.

Source: Wikipedia

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(Placozoa)

Placozoans

Пласт́инчасті

Placozoa is a phylum of free-living (non-parasitic) marine invertebrates. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by engulfment, reproducing by fission or budding, placozoans are described as “the simplest animals on Earth.” Structural and molecular analyses have supported them as among the most basal animals, thus, constituting a primitive metazoan phylum.

Placozoans do not have well-defined body plans, much like amoebas, unicellular eukaryotes. An individual body measures about 0.55 mm in diameter. There are no body parts. Animals studied in laboratories have bodies consisting of everything from hundreds to millions of cells.

Placozoans have only three anatomical parts as tissue layers inside its body: the upper, intermediate (middle) and lower epithelia. There are at least six different cell types. The upper epithelium is the thinnest portion and essentially comprises flat cells with their cell body hanging underneath the surface, and each cell having a cilium. Crystal cells are sparsely distributed near the marginal edge. A few cells have unusually large number of mitochondria. The middle layer is the thickest made up of numerous fiber cells, which contain mitochondrial complexes, vacuoles and endosymbiotic bacteria in the endoplasmic reticulum. The lower epithelium consists of numerous monociliated cylinder cells along with a few endocrine-like gland cells and lipophil cells. Each lipophil cell contains numerous middle-sized granules, one of which is a secretory granule.

The body axes of Hoilungia and Trichoplax are overtly similar to the oral–aboral axis of cnidarians, animals from another phylum with which they are most closely related. Structurally, they can not be distinguished from other placozoans, so that identification is purely on genetic (mitochondrial DNA) differences.

Trichoplax is a small, flattened, animal around 1 mm (0.039 in) across. An amorphous multi-celled body, analogous to a single-celled amoeba, it has no regular outline, although the lower surface is somewhat concave, and the upper surface is always flattened. The body consists of an outer layer of simple epithelium enclosing a loose sheet of stellate cells resembling the mesenchyme of some more complex animals. The epithelial cells bear cilia, which the animal uses to help it creep along the seafloor.

The lower surface engulfs small particles of organic detritus, on which the animal feeds. All placozoans can reproduce asexually, budding off smaller individuals, and the lower surface may also bud off eggs into the mesenchyme. Sexual reproduction has been reported to occur in one clade of placozoans, whose strain H8 was later found to belong to genus Cladtertia, where intergenic recombination was observed as well as other hallmarks of sexual reproduction.

Some Trichoplax species contain Rickettsiales bacteria as endosymbionts. One of the at least 20 described species turned out to have two bacterial endosymbionts; Grellia which lives in the animal’s endoplasmic reticulum and is assumed to play a role in the protein and membrane production. The other endosymbiont is the first described Margulisbacteria, that lives inside cells used for algal digestion. It appears to eat the fats and other lipids of the algae and provide its host with vitamins and amino acids in return.

Studies suggest that aragonite crystals in crystal cells have the same function as statoliths, allowing it to use gravity for spatial orientation.

Located in the dorsal epithelium there are lipid granules called shiny spheres which release a cocktail of venoms and toxins as an anti-predator defense, and can induce paralysis or death in some predators. Genes has been found in Trichoplax with a strong resemblance to the venom genes of some poisonous snakes, like the American copperhead and the West African carpet viper.

The Placozoa show substantial evolutionary radiation in regard to sodium channels, of which they have 5–7 different types, more than any other invertebrate species studied to date.

Three modes of population dynamics depended upon feeding sources, including induction of social behaviors, morphogenesis, and reproductive strategies.

In addition to fission, representatives of all species produced “swarmers” (a separate vegetative reproduction stage), which could also be formed from the lower epithelium with greater cell-type diversity.

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Trichoplax adhaerens

(Trichoplax adhaerens)

Trichoplax

Трихоплакс

Body length: 4mm.

Has been collected, among other places, in the Red Sea, the Mediterranean, and the Caribbean, off Hawaii, Guam, Samoa, Japan, Vietnam, Brazil, and Papua New Guinea, and on the Great Barrier Reef off the east coast of Australia.

Hoilungia hongkongensis

(Hoilungia hongkongensis)

Was discovered in brackish water from mangrove swamps in Hong Kong. These organisms are generally found in the biofilm surfaces in tropical and subtropical environments.

Cladtertia collaboinventa

(Cladtertia collaboinventa)

It has been found in warm tropical waters with low seasonal changes, ranging from 26°N to 25°S, where it occupies a distinct ecological niche compared to other placozoan lineages.

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(Dicyemida)

Dicyemids

Дицієміди

Dicyemida, also known as Rhombozoa, is a phylum of tiny parasites that live in the renal appendages of cephalopods. Adult dicyemids range in length from 0.5 to 7 mm, and they can be easily viewed through a light microscope. 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 ciliated 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. When more than one species of dicyemida exist within the same host, they have distinctly shaped calottes, which range in shape from conical to disk shaped, or cap shaped. To this day, there has never been a recorded case of two separate species of dicyemida existing in the same host and having exactly the same calotte.

Some part of the dicyemid life cycle may be tied to temperate benthic environments, where they occur in greatest abundance. While dicyemids have occasionally been found in the tropics, the infection rates are typically quite low, and many potential host species are not infected. Dicyemids have never been reported from truly oceanic cephalopods, who instead host a parasitic ciliate fauna.

The phylum contains three families, Conocyemidae, Dicyemidae and Kantharellidae.

Conocyemidae: Conocyema polymorpha, Conocyema deca, Microcyema vespa.

Dicyemidae: Dicyema microcephalum, Dicyema moschatum, Dicyema shorti, Dicyema acciaccatum, Dicyema acheroni, Dicyema aegira, Dicyema apalachiensis, Dicyema australis, Dicyema balamuthi, Dicyema banyulensis, Dicyema benedeni, Dicyema benthoctopi, Dicyema bilobum, Dicyema caudatum, Dicyema ganapatii, Dicyema colurum, Dicyema erythrum, Dicyema dolichocephalum. Dicyema sphyrocephalum, Dicyema lycidoeceum, Dicyema hadrum, Dicyema rhadinum, Dicyema hypercephalum, Dicyema robsonellae, Dicyema knoxi, Dicyema maorum, Dicyema madrasensis, Dicyema octopusi, Dicyema nouveli, Dicyema monodi, Dicyema oligomerum, Dicyema misakiense, Dicyema orientale, Dicyema plathycephalum, Dicyema typoides, Dicyema whitmani, Dicyema sullivani, Dicyema briarei, Dicyema megalocephalum, Dicyema typus, Dicyema acuticephalum, Dicyema apollyoni, Dicyema paradoxum, Dicyema japonicum, Dicyema clavatum, Dicyema schulzianum, Dicyema rondeletiolae, Dicyema macrocephalum, Pseudicyema nakaoi, Pseudicyema truncatum, Dicyemennea gyrinodes, Dicyemennea ophioides, Dicyemennea trochocephalum, Dicyemennea mastigoides, Dicyemennea minabense, Dicyemennea granularis, Dicyemennea abelis, Dicyemennea adscita, Dicyemennea abasi, Dicyemennea abbreviata, Dicyemennea bathybenthum, Dicyemennea lameerei, Dicyemennea antarcticensis, Dicyemennea abreida, Dicyemennea brevicephala, Dicyemennea californica, Dicyemennea filiformis, Dicyemennea rossiae, Dicyemennea brevicephaloides, Dicyemennea longinucleata, Dicyemennea longinucleata, Dicyemennea kaikouriensis, Dicyemennea rostrata, Dicyemennea eltanini, Dicyemennea littlei, Dicyemennea discocephala, Dicyemennea coromandelensis, Dicyemennea eledones, Dicyemennea gracile, Dicyemennea dogieli, Dicyemennea dorycephalum, Dicyemennea canadensis, Dicyemennea adminicula, Dicyemennea parva, Dicyemennea marplatensis, Dicyemennea nouveli, Dicyemodeca anthinocephalum, Dicyemodeca sceptrum, Dicyemodeca dogieli, Pleodicyema delamarie, Dodecadicyema loligoi.

Katharellidae: Kantharella antarctica

Dicyema Japonicum
(Dicyema japonicum)
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(Orthonectida)

Orthonectids

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Orthonectida is a small phylum of poorly-known parasites of marine invertebrates. They are among the simplest of multi-cellular organisms.

The adults are microscopic wormlike animals. They have a single layer of outer cells surrounding a mass of sex cells. They swim freely within the bodies of their hosts, which include flatworms, polychaete worms, bivalve molluscs, and echinoderms. They have separate male and female individuals.

When they are ready to reproduce, the adults are released from the host, and sperm from the males penetrates the bodies of the females to achieve fertilisation. The resulting zygote develops into a ciliated larva that escapes from the mother to seek out new hosts. Once it finds a host, the larva loses its cilia and develops into a syncytial plasmodium larva. This, in turn, breaks up into numerous individual cells that become the next generation of adults.

About 30 species of orthonectids are known, united into five genera and two families: Rhopaluridae, Pelmatosphaeridae.

Rhopaluridae: Ciliocincta sabellariae, Ciliocincta akkeshiensis, Ciliocincta julini, Intoshia linei, Intoshia leptoplanae, Intoshia paraphanostomae, Rhopalura elongata, Rhopalura granosa, Rhopalura intoshi, Rhopalura philinae, Rhopalura ophiocomae, Rhopalura murmanica, Rhopalura litoralis, Rhopalura major, Intoshia metchnikovi, Stoecharthrum giardi, Rhopalura pterocirri, Rhopalura pelseneeri, Rhopalura vermiculicola, Rhopalura sanguinea, Stoecharthrum monnati, Stoecharthrum fosterae, Stoecharthrum burresoni, Intoshia variabili.

Pelmatosphaeridae: Pelmatosphaera polycirri.

Intoshia linei
(Intoshia linei)
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