Echinoderm

Last updated

Echinoderms
Temporal range: Cambrian–recent
Є
O
S
D
C
P
T
J
K
Pg
N
Brockhaus and Efron Encyclopedic Dictionary b24 782-0.jpg
Various echinoderms
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Xenambulacraria
Clade: Ambulacraria
Phylum:Echinodermata
Bruguière, 1791 [ex Klein, 1734]
Subphyla and classes [1]

HomalozoaGill & Caster, 1960

Homostelea
Homoiostelea
Stylophora
CtenocystoideaRobison & Sprinkle, 1969

Crinozoa

Crinoidea
Edrioasteroidea
? Arkarua
Cystoidea
Rhombifera

Asterozoa

Ophiuroidea
Asteroidea

Echinozoa

Echinoidea
Holothuroidea
Ophiocistioidea
Helicoplacoidea

Blastozoa

Blastoidea
Cystoideavon Buch, 1846
Eocrinoidea Jaekel, 1899
ParacrinoideaRegnéll, 1945

† = Extinct

Echinoderm is the common name given to any member of the phylum Echinodermata (from Ancient Greek, ἐχῖνος, echinos – "hedgehog" and δέρμα, derma – "skin") [2] of marine animals. The adults are recognizable by their (usually five-point) radial symmetry, and include such well-known animals as starfish, sea urchins, sand dollars, and sea cucumbers, as well as the sea lilies or "stone lilies". [3] Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 7000 living species, [4] making it the second-largest grouping of deuterostomes (a superphylum), after the chordates (which include the vertebrates, such as birds, fishes, mammals, and reptiles). Echinoderms are also the largest phylum that has no freshwater or terrestrial (land-based) representatives.

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia or Metazoa contains approximately 35 phyla, the plant kingdom Plantae contains about 14, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Ancient Greek Version of the Greek language used from roughly the 9th century BCE to the 6th century CE

The ancient Greek language includes the forms of Greek used in Ancient Greece and the ancient world from around the 9th century BCE to the 6th century CE. It is often roughly divided into the Archaic period, Classical period, and Hellenistic period. It is antedated in the second millennium BCE by Mycenaean Greek and succeeded by Medieval Greek.

Animal Kingdom of motile multicellular eukaryotic heterotrophic organisms

Animals are multicellular eukaryotic organisms that form the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Over 1.5 million living animal species have been described—of which around 1 million are insects—but it has been estimated there are over 7 million animal species in total. Animals range in length from 8.5 millionths of a metre to 33.6 metres (110 ft). They have complex interactions with each other and their environments, forming intricate food webs. The kingdom Animalia includes humans, but in colloquial use the term animal often refers only to non-human animals. The study of non-human animals is known as zoology.

Contents

Aside from the hard-to-classify Arkarua (a Precambrian animal with echinoderm-like pentamerous radial symmetry), the first definitive members of the phylum appeared near the start of the Cambrian. One group of Cambrian echinoderms, the cinctans (Homalozoa), which are close to the base of the echinoderm origin, have been found to possess external gills used for filter feeding, similar to those possessed by chordates and hemichordates. [5]

<i>Arkarua</i>

Arkarua adami is a small, Precambrian disk-like fossil with a raised center, a number of radial ridges on the rim, and a five-pointed central depression marked with radial lines of 5 small dots from the middle of the disk center. Fossils range from 3 to 10 mm in diameter.

The Precambrian is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of the Phanerozoic eon, which is named after Cambria, the Latinised name for Wales, where rocks from this age were first studied. The Precambrian accounts for 88% of the Earth's geologic time.

The Cambrian Period was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 55.6 million years from the end of the preceding Ediacaran Period 541 million years ago (mya) to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, and its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name of Wales, where Britain's Cambrian rocks are best exposed. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. As a result, our understanding of the Cambrian biology surpasses that of some later periods.

The echinoderms are important both ecologically and geologically. Ecologically, there are few other groupings so abundant in the biotic desert of the deep sea, as well as shallower oceans. Most echinoderms are able to reproduce asexually and regenerate tissue, organs, and limbs; in some cases, they can undergo complete regeneration from a single limb. Geologically, the value of echinoderms is in their ossified skeletons, which are major contributors to many limestone formations, and can provide valuable clues as to the geological environment. They were the most used species in regenerative research in the 19th and 20th centuries. Further, it is held by some scientists that the radiation of echinoderms was responsible for the Mesozoic Marine Revolution.

Continental shelf A portion of a continent that is submerged under an area of relatively shallow water known as a shelf sea

A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water known as a shelf sea. Much of the shelves were exposed during glacial periods and interglacial periods. The shelf surrounding an island is known as an insular shelf.

The deep sea or deep layer is the lowest layer in the ocean, existing below the thermocline and above the seabed, at a depth of 1000 fathoms or more. Little or no light penetrates this part of the ocean, and most of the organisms that live there rely for subsistence on falling organic matter produced in the photic zone. For this reason, scientists once assumed that life would be sparse in the deep ocean, but virtually every probe has revealed that, on the contrary, life is abundant in the deep ocean.

From the time of Pliny until the late nineteenth century...humans believed there was no life in the deep. It took a historic expedition in the ship Challenger between 1872 and 1876 to prove Pliny wrong; its deep-sea dredges and trawls brought up living things from all depths that could be reached. Yet even in the twentieth century scientists continued to imagine that life at great depth was insubstantial, or somehow inconsequential. The eternal dark, the almost inconceivable pressure, and the extreme cold that exist below one thousand meters were, they thought, so forbidding as to have all but extinguished life. The reverse is in fact true....(Below 200 meters) lies the largest habitat on earth.

Asexual reproduction Biological process in which new individuals are produced by either a single cell or a group of cells, in the absence of any sexual process

Asexual reproduction is a type of reproduction by which offspring arise from a single organism, and inherit the genes of that parent only; it does not involve the fusion of gametes, and almost never changes the number of chromosomes. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many plants and fungi sometimes reproduce asexually. Some asexual cells die when they are very young.

Taxonomy and evolution

Early echinoderms EarlyEchinoderms NT.jpg
Early echinoderms

Along with the chordates and hemichordates, echinoderms are deuterostomes, one of the two major divisions of the bilaterians, the other being the protostomes. During the early development of the embryo, in deuterostomes, the blastopore (the first opening to form) becomes the anus whereas in the protostomes, it becomes the mouth. In deuterostomes, the mouth develops at a later stage, at the opposite end of the blastula from the blastopore, and a gut forms connecting the two. [6] The larvae of echinoderms have bilateral symmetry but this is lost during metamorphosis when their bodies are reorganised and develop the characteristic radial symmetry of the echinoderm, typically pentamerism. [7] The characteristics of adult echinoderms are the possession of a water vascular system with external tube feet and a calcareous endoskeleton consisting of ossicles connected by a mesh of collagen fibres. [8] A 2014 analysis of 219 genes from all classes of echinoderms gives the following phylogenetic tree. [9]

Deuterostome Superphylum of bilateral animals

Deuterostomes comprise a superphylum of animals. It is a sister clade of Protostomia, with which it forms the Nephrozoa clade.

Bilateria all animals having a bilateral symmetry

The bilateria, bilaterians, or triploblasts, are animals with bilateral symmetry, i.e., they have a head (anterior) and a tail (posterior) as well as a back (dorsal) and a belly (ventral); therefore they also have a left side and a right side.

Protostome Clade of animals

Protostomia is a clade of animals. Together with the deuterostomes and xenacoelomorpha, its members make up the Bilateria, mostly comprising animals with bilateral symmetry and three germ layers. The major distinctions between deuterostomes and protostomes are found in embryonic development and is based on the embryological origins of the mouth and anus.

Bilateria

Xenacoelomorpha Proporus sp.png

Nephrozoa
Deuterostomia

Chordata and allies Cyprinus carpio3.jpg

Echinodermata
Echinozoa

Holothuroidea Holothuroidea.JPG

Echinoidea S. variolaris.jpg

Asterozoa

Ophiuroidea Ophiura ophiura.jpg

Asteroidea Portugal 20140812-DSC01434 (21371237591).jpg

Crinoidea Crinoid on the reef of Batu Moncho Island.JPG

Protostomia

Ecdysozoa Long nosed weevil edit.jpg

Spiralia Grapevinesnail 01.jpg

610 mya
650 mya
The Ordovician cystoid Echinosphaerites from northeastern Estonia Echinosphaerites.JPG
The Ordovician cystoid Echinosphaerites from northeastern Estonia

There are a total of about 7,000 extant species of echinoderm as well as about 13,000 extinct species. [8] They are found in habitats ranging from shallow intertidal areas to abyssal depths. Two main subdivisions are traditionally recognised: the more familiar motile Eleutherozoa, which encompasses the Asteroidea (starfish, 1,745 recent species), Ophiuroidea (brittle stars, 2,300 species), Echinoidea (sea urchins and sand dollars, 900 species) and Holothuroidea (sea cucumbers, 1,430 species); and the Pelmatozoa, some of which are sessile while others move around. These consist of the Crinoidea (feather stars and sea lilies, 580 species) and the extinct blastoids and Paracrinoids. [10] A fifth class of Eleutherozoa consisting of just three species, the Concentricycloidea (sea daisies), were recently merged into the Asteroidea. [11] The fossil record includes a large number of other classes which do not appear to fall into any extant crown group.

Motility Ability to move spontaneously and actively, using metabolic energy

Motility is the ability of an organism to move independently, using metabolic energy. This is in contrast to mobility, which describes the ability of an object to be moved. Motility is genetically determined, but may be affected by environmental factors. For instance, muscles give animals motility but the consumption of hydrogen cyanide would adversely affect muscle physiology, causing them to stiffen, leading to rigor mortis. In addition to animal locomotion, most animals are motile. The term applies to bacteria and other microorganisms, and to some multicellular organisms, as well as to some mechanisms of fluid flow in multicellular organs and tissue. Motile marine animals are commonly called free-swimming, and motile non-parasitic organisms are called free-living.

Eleutherozoa subphylum of echinoderms

Eleutherozoa is a proposed subphylum of echinoderms. They are mobile animals with the mouth directed towards the substrate. They usually have a madreporite, tube feet, and moveable spines of some sort, and some have Tiedemann's bodies on the ring canal. All living echinoderms except Crinoidea belong here.

Starfish class of echinoderms, marine animal

Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea. Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. About 1,500 species of starfish occur on the seabed in all the world's oceans, from the tropics to frigid polar waters. They are found from the intertidal zone down to abyssal depths, 6,000 m (20,000 ft) below the surface.

Fossil crinoid crowns Fossile-seelilie.jpg
Fossil crinoid crowns

All echinoderms are marine and nearly all are benthic. [12] The oldest known echinoderm fossil may be Arkarua from the Precambrian of Australia. It is a disc-like fossil with radial ridges on the rim and a five-pointed central depression marked with radial lines. However, no stereom or internal structure showing a water vascular system is present and the identification is inconclusive. [13]

Marine biology The scientific study of organisms that live in the ocean

Marine biology is the scientific study of marine life, organisms in the sea. Given that in biology many phyla, families and genera have some species that live in the sea and others that live on land, marine biology classifies species based on the environment rather than on taxonomy.

Fossil Preserved remains or traces of organisms from a past geological age

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood, oil, coal, and DNA remnants. The totality of fossils is known as the fossil record.

Stereom is a calcium carbonate material that makes up the internal skeletons found in sea urchins, and all other echinoderms, both living and fossilized forms. It is a sponge-like porous structure which, in a sea urchin may be 50% by volume living cells, and the rest being a matrix of calcite crystals. The size of openings in stereom varies in different species and in different places within the same organism. When an echinoderm becomes a fossil, microscopic examination is used to reveal the structure and such examination is often an important tool to classify the fossil as an echinoderm or related creature.

The first universally accepted echinoderms appear in the Lower Cambrian period, asterozoans appeared in the Ordovician and the crinoids were a dominant group in the Paleozoic. [12] Echinoderms left behind an extensive fossil record. [12] It is hypothesised that the ancestor of all echinoderms was a simple, motile, bilaterally symmetrical animal with a mouth, gut and anus. This ancestral stock adopted an attached mode of life and suspension feeding, and developed radial symmetry as this was more advantageous for such an existence. The larvae of all echinoderms are even now bilaterally symmetrical and all develop radial symmetry at metamorphosis. The starfish and crinoids still attach themselves to the seabed while changing to their adult form. [14]

The first echinoderms later gave rise to free-moving groups. The evolution of endoskeletal plates with stereom structure and of external ciliary grooves for feeding were early echinoderm developments. [15] The Paleozoic echinoderms were globular, attached to the substrate and were orientated with their oral surfaces upwards. The fossil echinoderms had ambulacral grooves extending down the side of the body, fringed on either side by brachioles, structures very similar to the pinnules of a modern crinoid. It seems probable that the mouth-upward orientation is the primitive state and that at some stage, all the classes of echinoderms except the crinoids reversed this to become mouth-downward. Before this happened, the podia probably had a feeding function as they do in the crinoids today. Their locomotor function came later, after the re-orientation of the mouth when the podia were in contact with the substrate for the first time. [14]

Anatomy and physiology

Echinoderms evolved from animals with bilateral symmetry. Although adult echinoderms possess pentaradial, or five-sided, symmetry, echinoderm larvae are ciliated, free-swimming organisms that organize in bilateral symmetry which makes them look like embryonic chordates. Later, the left side of the body grows at the expense of the right side, which is eventually absorbed. The left side then grows in a pentaradially symmetric fashion, in which the body is arranged in five parts around a central axis. [16] Within the Asterozoa, there can be a few exceptions from the rule. The starfish genus Leptasterias normally have six arms, although five-armed individuals can occur. Also the Brisingida have six armed species. Amongst the brittle stars, six-armed species such as Ophiothela danae, Ophiactis savignyi and Ophionotus hexactis exists, and Ophiacantha vivipara often has more than six. [17]

Echinoderms exhibit secondary radial symmetry in portions of their body at some stage of life. This, however, is an adaptation to their sessile existence. They developed from other members of the Bilateria and exhibit bilateral symmetry in their larval stage. Many crinoids and some seastars exhibit symmetry in multiples of the basic five, with starfish such as Labidiaster annulatus known to possess up to fifty arms, and the sea-lily Comaster schlegelii having two hundred. [18]

Skin and skeleton

A brittle star, Ophionereis reticulata Reef2589.jpg
A brittle star, Ophionereis reticulata
A sea cucumber from Malaysia Sea cucumber at Pulau Redang.jpg
A sea cucumber from Malaysia
Starfish exhibit a wide range of colours Nerr0878.jpg
Starfish exhibit a wide range of colours
Strongylocentrotus purpuratus, a well-armoured sea urchin Strongylocentrotus purpuratus 1.jpg
Strongylocentrotus purpuratus , a well-armoured sea urchin
Crinoid on a coral reef Crinoid on the reef of Batu Moncho Island.JPG
Crinoid on a coral reef

Echinoderms have a mesodermal skeleton composed of calcareous plates or ossicles. Each one of these, even the articulating spine of a sea urchin, is composed mineralogically of a crystal of calcite. If solid, these would form a heavy skeleton, so they have a sponge-like porous structure known as stereom. [19] Ossicles may be fused together, as in the test of sea urchins, or may articulate with each other as in the arms of sea stars, brittle stars and crinoids. The ossicles may be flat plates or bear external projections in the form of spines, granules or warts and they are supported by a tough epidermis (skin). Skeletal elements are also deployed in some specialized ways, such as the "Aristotle's lantern" mouthparts of sea urchins used for grinding, the supportive stalks of crinoids and the structural "lime ring" of sea cucumbers. [16]

Despite the robustness of the individual skeletal modules complete skeletons of starfish, brittle stars and crinoids are rare in the fossil record. This is because they quickly disarticulate (disconnect from each other) once the encompassing skin rots away, and in the absence of tissue there is nothing to hold the plates together. The modular construction is a result of the growth system employed by echinoderms, which adds new segments at the centre of the radial limbs, pushing the existing plates outwards and lengthening the arms. Sea urchins on the other hand are often well preserved in chalk beds or limestone. During fossilization, the cavities in the stereom are filled in with calcite that is in crystalline continuity with the surrounding material. On fracturing such rock, distinctive cleavage patterns can be seen and sometimes even the intricate internal and external structure of the test. [20]

The epidermis consists of cells responsible for the support and maintenance of the skeleton, as well as pigment cells, mechanoreceptor cells (which detect motion on the animal's surface), and sometimes gland cells which secrete sticky fluids or even toxins. The varied and often vivid colours of echinoderms are produced by the action of skin pigment cells. These are produced by a variable combination of coloured pigments, such as the dark melanin, red carotinoids, and carotene proteins, which can be blue, green, or violet. These may be light-sensitive, and as a result many echinoderms change appearance completely as night falls. The reaction can happen quickly — the sea urchin Centrostephanus longispinus changes from jet black to grey-brown in just fifty minutes when exposed to light. [21]

One characteristic of most echinoderms is a special kind of tissue known as "catch connective tissue". This collagenous material can change its mechanical properties in a few seconds or minutes through nervous control rather than by muscular means. This tissue enables a starfish to change from moving flexibly around the seabed to becoming rigid while prying open a bivalve mollusc or preventing itself from being extracted from a crevice. Similarly, sea urchins can lock their normally mobile spines rigidly as a defensive mechanism when attacked. [22]

The water vascular system

Echinoderms possess a unique water vascular system. This is a network of fluid-filled canals derived from the coelom (body cavity) that function in gas exchange, feeding, sensory reception and locomotion. This system varies between different classes of echinoderm but typically opens to the exterior through a sieve-like madreporite on the aboral (upper) surface of the animal. The madreporite is linked to a slender duct, the stone canal, which extends to a ring canal that encircles the mouth or oesophagus. From this, radial canals extend along the arms of asteroids and adjoin the test in the ambulacral areas of echinoids. Short lateral canals branch off the radial canals, each one ending in an ampulla. Part of the ampulla can protrude through a pore (or a pair of pores in sea urchins) to the exterior and is known as a podium or tube feet. The water vascular system assists with the distribution of nutrients throughout the animal's body and is most obviously expressed in the tube feet which can be extended or contracted by the redistribution of fluid between the foot and the internal sac. [23]

The organization of the system is somewhat different in ophiuroids where the madreporite may be on the oral surface and the podia lack suckers. [24] In holothuroids, the podia may be reduced or absent and the madreporite opens into the body cavity so that the circulating liquid is coelomic fluid rather than sea water. [25] The arrangements in crinoids is similar to asteroids but the tube feet lack suckers and are used to pass food particles captured by the arms towards the central mouth. In the asteroids, the same wafting motion is employed to move the animal across the ground. [26] Sea urchins use their feet to prevent the larvae of encrusting organisms from settling on their surfaces; potential settlers are moved to the urchin's mouth and eaten. Some burrowing sea stars extend their elongated dorsal tube feet to the surface of the sand or mud above and use them to absorb oxygen from the water column. [27]

Other organs

Echinoderms possess a simple digestive system which varies according to the animal's diet. Starfish are mostly carnivorous and have a mouth, oesophagus, two-part stomach, intestine and rectum, with the anus located in the centre of the aboral body surface. With a few exceptions, the members of the order Paxillosida do not possess an anus. [28] [29] In many species of starfish, the large cardiac stomach can be everted and digest food outside the body. In other species, whole food items such as molluscs may be ingested. [30] Brittle stars have a blind gut with no intestine or anus. They have varying diets and expel food waste through their mouth. [31] Sea urchins are herbivores and use their specialised mouthparts to graze, tear and chew algae and sometimes other animal or vegetable material. They have an oesophagus, a large stomach and a rectum with the anus at the apex of the test. [32] Sea cucumbers are mostly detritivores, sorting through the sediment with their buccal tentacles which are modified tube feet. Sand and mud accompanies their food through their simple gut which has a long coiled intestine and a capacious cloaca. [33] Crinoids are passive suspension feeders, catching plankton with their outstretched arms. Boluses of mucus-trapped food are passed to the mouth which is linked to the anus by a loop consisting of a short oesophagus and longer intestine. [34]

The coelomic cavities of echinoderms are complex. Aside from the water vascular system, echinoderms have a haemal coelom (or haemal system, the "haemal" being a misnomer), a perivisceral coelom, a gonadal coelom and often also a perihaemal coelom (or perihaemal system). [35] During development, echinoderm coelom is divided in metacoel, mesocoel and protocoel (also called somatocoel, hydrocoel and axocoel, respectively). [36] The water vascular system, haemal system and perihaemal system form the tubular coelomic system. [37] Echinoderms are an exception having both a coelomic circulatory system (i.e., the water vascular system) and a haemal circulatory system (i.e., the haemal and perihaemal systems). [38]

Haemal and perihaemal systems are derived from the coelom and form an open and reduced circulatory system. This usually consists of a central ring and five radial vessels. There is no true heart and the blood often lacks any respiratory pigment. Gaseous exchange occurs via dermal branchae or papulae in starfish, genital bursae in brittle stars, peristominal gills in sea urchins and cloacal trees in sea cucumbers. Exchange of gases also takes place through the tube feet. Echinoderms lack specialized excretory (waste disposal) organs and so nitrogenous waste, chiefly in the form of ammonia, diffuses out through the respiratory surfaces. [23]

The coelomic fluid contains the coelomocytes, or immune cells. There are several types of immune cells, which vary among classes and species. All classes possess a type of phagocytic amebocyte, which engulf invading particles and infected cells, aggregate or clot, and may be involved in cytotoxicity. These cells are usually larger and granular, and are suggested to be a main line of defense against potential pathogens. [39] Depending on the class, echinoderms may have spherule cells (for cytotoxicity, inflammation, and anti-bacterial activity), vibratile cells (for coelomic fluid movement and clotting), and crystal cells (potential osmoregulatory cells in sea cucumbers),. [39] [40] The coelomocytes also secrete Anti-Microbial Peptides (AMPs) against bacteria, and have a set of lectins and complement proteins as part of an innate immune system that is still being characterized. [2]

Echinoderms have a simple radial nervous system that consists of a modified nerve net consisting of interconnecting neurons with no central brain, although some do possess ganglia. Nerves radiate from central rings around the mouth into each arm or along the body wall; the branches of these nerves coordinate the movements of the organism and the synchronisation of the tube feet. Starfish have sensory cells in the epithelium and have simple eyespots and touch-sensitive tentacle-like tube feet at the tips of their arms. Sea urchins have no particular sense organs but do have statocysts that assist in gravitational orientation, and they have sensory cells in their epidermis, particularly in the tube feet, spines and pedicellariae. Brittle stars, crinoids and sea cucumbers in general do not have sensory organs but some burrowing sea cucumbers of the order Apodida have a single statocyst adjoining each radial nerve and some have an eyespot at the base of each tentacle. [41]

The gonads occupy much of the body cavities of sea urchins and sea cucumbers, while the less voluminous crinoids, brittle stars and starfish have two gonads in each arm. While the ancestral condition is considered to be the possession of one genital aperture, many organisms have multiple gonopores through which eggs or sperm may be released. [41]

Regeneration

Sunflower star regenerating several arms Sea star regenerating legs.jpg
Sunflower star regenerating several arms

Many echinoderms have remarkable powers of regeneration. Many species routinely autotomize and regenerate arms and viscera. Sea cucumbers often discharge parts of their internal organs if they perceive themselves to be threatened. The discharged organs and tissues are regenerated over the course of several months. Sea urchins are constantly replacing spines lost through damage. Sea stars and sea lilies readily lose and regenerate their arms. In most cases, a single severed arm cannot grow into a new starfish in the absence of at least part of the disc. [42] [43] [44] [45] However, in a few species a single arm can survive and develop into a complete individual [43] [44] [45] and in some species, the arms are intentionally detached for the purpose of asexual reproduction. During periods when they have lost their digestive tracts, sea cucumbers live off stored nutrients and absorb dissolved organic matter directly from the water. [46]

The regeneration of lost parts involves both epimorphosis and morphallaxis. In epimorphosis stem cells—either from a reserve pool or those produced by dedifferentiation—form a blastema and generate new tissues. Morphallactic regeneration involves the movement and remodelling of existing tissues to replace lost parts. Direct transdifferentiation of one type of tissue to another during tissue replacement is also observed. [47]

The robust larval growth is responsible for many echinoderms being used as popular model organisms in developmental biology. [48]

Reproduction

Sexual reproduction

Echinoderms become sexually mature after approximately two to three years, depending on the species and the environmental conditions. They are nearly all gonochoric, though a few species are hermaphroditic. The eggs and sperm cells are typically released into open water, where fertilization takes place. The release of sperm and eggs is synchronised in some species, usually with regard to the lunar cycle. In other species, individuals may aggregate during the reproductive season, thereby increasing the likelihood of successful fertilisation. Internal fertilisation has currently been observed in three species of sea star, three brittle stars and a deep water sea cucumber. Even at abyssal depths, where no light penetrates, synchronisation of reproductive activity in echinoderms is surprisingly frequent. [49]

Some echinoderms brood their eggs. This is especially common in cold water species where planktonic larvae might not be able to find sufficient food. These retained eggs are usually few in number and are supplied with large yolks to nourish the developing embryos. In starfish, the female may carry the eggs in special pouches, under her arms, under her arched body or even in her cardiac stomach. [50] Many brittle stars are hermaphrodites. Egg brooding is quite common and usually takes place in special chambers on their oral surfaces, but sometimes the ovary or coelom is used. [51] In these starfish and brittle stars, direct development without passing through a bilateral larval stage usually takes place. [52] A few sea urchins and one species of sand dollar carry their eggs in cavities, or near their anus, holding them in place with their spines. [53] Some sea cucumbers use their buccal tentacles to transfer their eggs to their underside or back where they are retained. In a very small number of species, the eggs are retained in the coelom where they develop viviparously, later emerging through ruptures in the body wall. [54] In some species of crinoid, the embryos develop in special breeding bags, where the eggs are held until sperm released by a male happens to find them. [55]

Asexual reproduction

One species of seastar, Ophidiaster granifer , reproduces asexually by parthenogenesis. [56] In certain other asterozoans, the adults reproduce asexually for a while before they mature after which time they reproduce sexually. In most of these species, asexual reproduction is by transverse fission with the disc splitting in two. Regrowth of both the lost disc area and the missing arms occur [45] [57] so that an individual may have arms of varying lengths. Though in most species at least part of the disc is needed for complete regeneration, in a few species of sea stars, a single severed arm can grow into a complete individual over a period of several months. [43] [44] [45] In at least some of these species, they actively use this as a method of asexual reproduction. [43] [58] A fracture develops on the lower surface of the arm and the arm pulls itself free from the body which holds onto the substrate during the process. [58] During the period of regrowth, they have a few tiny arms and one large arm, thus often being referred to as "comets". [44] [58]

Asexual reproduction by transverse fission has also been observed in adult sea cucumbers. Holothuria parvula uses this method frequently, an individual splitting into two a little in front of the midpoint. The two halves each regenerate their missing organs over a period of several months but the missing genital organs are often very slow to develop. [59]

The larvae of some echinoderm species are capable of asexual reproduction. This has long been known to occur among starfish and brittle stars but has been more recently observed in a sea cucumber, a sand dollar and a sea urchin. These species belong to four of the major classes of echinoderms except crinozoans (as of 2011). [60] Asexual reproduction in the planktonic larvae occurs through numerous modes. They may autotomise parts that develop into secondary larvae, grow buds or undergo paratomy. The parts that are autotomised or the buds may develop directly into fully formed larvae or may develop through a gastrula or even a blastula stage. The parts that develop into the new larvae vary from the preoral hood (a mound like structure above the mouth), the side body wall, the postero-lateral arms or their rear ends. [60] [61] [62]

The process of cloning is a cost borne by the larva both in resources as well as in development time. Larvae have been observed to undergo this process when food is plentiful [63] or temperature conditions are optimal. [62] It has also been suggested that cloning may occur to make use of the tissues that are normally lost during metamorphosis. [64] Recent research has shown that the larvae of some sand dollars clone themselves when they detect predators (by sensing dissolved fish mucus). [62] [64] Asexual reproduction produces many smaller larvae that escape better from planktivorous fish. [65]

Larval development

An echinopluteus larva with larval arms Pluteus001.jpg
An echinopluteus larva with larval arms

The development of an echinoderm begins with a bilaterally symmetrical embryo, with a coeloblastula developing first. Gastrulation marks the opening of the "second mouth" that places echinoderms within the deuterostomes, and the mesoderm, which will host the skeleton, migrates inwards. The secondary body cavity, the coelom, forms by the partitioning of three body cavities. The larvae are mostly planktonic but in some species the eggs are retained inside the female and in some, the larvae are also brooded by the female. [66]

The larvae of echinoderms pass through a number of stages and these have specific names derived from the taxonomic names of the adults or from their appearance. For example, a sea urchin has an 'echinopluteus' larva while a brittle star has an 'ophiopluteus' larva. A starfish has a 'bipinnaria' larva but this later develops into a multi-armed 'brachiolaria' larva. A sea cucumber larva is an 'auricularia' while a crinoid one is a 'vitellaria'. All these larvae are bilaterally symmetrical and have bands of cilia with which they swim and some, usually known as 'pluteus' larvae, have arms. When fully developed they settle on the seabed to undergo metamorphosis and the larval arms and gut degenerate. The left hand side of the larva develops into the oral surface of the juvenile while the right side becomes the aboral surface. At this stage the bilateral symmetry is lost and radial symmetry develops. [66] [67]

The planktotrophic larva is considered to be the ancestral larval type for echinoderms but after 500 million years of larval evolution, about 68% of species whose development is known have a lecithotrophic larval type. [10] The provision of a yolk-sac means that smaller numbers of eggs are produced, the larvae have a shorter development period, smaller dispersal potential but a greater chance of survival. There seems to be an evolutionary trend towards a "lower-risk–lower-gain" strategy of direct development. [10]

Distribution and habitat

Echinoderms are globally distributed in almost all depths, latitudes and environments in the ocean. They reach highest diversity in reef environments but are also widespread on shallow shores, around the poles — refugia where crinoids are at their most abundant — and throughout the deep ocean, where bottom-dwelling and burrowing sea cucumbers are common — sometimes accounting for up to 90% of organisms. While almost all echinoderms are benthic — that is, they live on the sea floor — some sea-lilies can swim at great velocity for brief periods of time, and a few deep-sea sea cucumbers are fully floating. Some crinoids are pseudo-planktonic, attaching themselves to floating logs and debris, although this behaviour was exercised most extensively in the Paleozoic, before competition from such organisms as barnacles restricted the extent of the behaviour. [68]

The larvae of echinoderms, especially starfish and sea urchins, are pelagic, and with the aid of ocean currents can be transported for great distances, reinforcing the global distribution of the phylum. [69]

Mode of life

Locomotion

Echinoderms primarily use their tube feet to move about but some sea urchins also use their spines. The tube feet typically have a tip shaped like a suction pad in which a vacuum can be created by contraction of muscles. This along with some stickiness provided by the secretion of mucus provides adhesion. Waves of tube feet contractions and relaxations move along the adherent surface and the animal moves slowly along. [70]

Brittle stars are the most agile of the echinoderms, raising their discs and taking strides when moving. The two forward arms grip the substrate with their tube feet, the two side arms "row", the hindermost arm trails and the animal moves in jerks. The arm spines provide traction and when moving among objects, the supple arms can coil around things. A few species creep around on pointed tube feet. [70] Starfish extend their tube feet in the intended direction of travel and grip the substrate by suction, after which the feet are drawn backwards. The movement of multiple tube feet, coordinated in waves, moves the animal forward, but progress is slow. [71] Some burrowing starfish have points rather than suckers on their tube feet and they are able to "glide" across the seabed at a faster rate. [72]

Sea urchins use their tube feet to move around in a similar way to starfish. Some also use their articulated spines to push or lever themselves along or lift their oral surfaces off the substrate. If a sea urchin is overturned, it can extend its tube feet in one ambulacral area far enough to bring them within reach of the substrate and then successively attach feet from the adjoining area until it is righted. Some species bore into rock and they usually do this by grinding away at the surface with their mouthparts. [73]

Sea cucumbers are generally sluggish animals. Many can move on the surface or burrow through sand or mud using peristaltic movements and some have short tube feet on their under surface with which they can creep along in the manner of a starfish. Some species drag themselves along by means of their buccal tentacles while others can expand and contract their body or rhythmically flex it and "swim". Many live in cracks, hollows and burrows and hardly move at all. Some deep water species are pelagic and can float in the water with webbed papillae forming sails or fins. [74]

The majority of crinoids are motile but the sea lilies are sessile and attached to hard substrates by stalks. These stems can bend and the arms can roll and unroll and that is about the limit of the sea lily's movement, although a few species can relocate themselves on the seabed by crawling. The sea feathers are unattached and usually live in crevices, under corals or inside sponges with their arms the only visible part. Some sea feathers emerge at night and perch themselves on nearby eminences to better exploit the food-bearing current. Many species can "walk" across the seabed, raising their body with the help of their arms. Many can also swim with their arms but most are largely sedentary, seldom moving far from their chosen place of concealment. [75]

Feeding

The modes of feeding vary greatly between the different echinoderm taxa. Crinoids and some brittle stars tend to be passive filter-feeders, enmeshing suspended particles from passing water; most sea urchins are grazers, sea cucumbers deposit feeders and the majority of starfish are active hunters.

Crinoids are suspension feeders and spread their arms wide to catch particles floating past. These are caught by the tube feet on the pinnules, moved into the ambulacral grooves, wrapped in mucus and conveyed to the mouth by the cilia lining the grooves. [76] The exact dietary requirements of crinoids have been little researched but in the laboratory they can be fed with diatoms. [77]

Basket stars are suspension feeders, raising their branched arms to collect zooplankton, while brittle stars use several methods of feeding, though usually one predominates. Some are suspension feeders, securing food particles with mucus strands, spines or tube feet on their raised arms. Others are scavengers and feeders on detritus. Others again are voracious carnivores and able to lasso their waterborne prey with a sudden encirclement by their flexible arms. The limbs then bend under the disc to transfer the food to the jaws and mouth. [78]

Many sea urchins feed on algae, often scraping off the thin layer of algae covering the surfaces of rocks with their specialised mouthparts known as Aristotle's lantern. Other species devour smaller organisms, which they may catch with their tube feet. They may also feed on dead fish and other animal matter. [79] Sand dollars may perform suspension feeding and feed on phytoplankton, detritus, algal pieces and the bacterial layer surrounding grains of sand. [80]

Many sea cucumbers are mobile deposit or suspension feeders, using their buccal podia to actively capture food and then stuffing the particles individually into their buccal cavities. Others ingest large quantities of sediment, absorb the organic matter and pass the indigestible mineral particles through their guts. In this way they disturb and process large volumes of substrate, often leaving characteristic ridges of sediment on the seabed. Some sea cucumbers live infaunally in burrows, anterior-end down and anus on the surface, swallowing sediment and passing it through their gut. Other burrowers live anterior-end up and wait for detritus to fall into the entrances of the burrows or rake in debris from the surface nearby with their buccal podia. [81]

Nearly all starfish are detritivores or carnivores, though a few are suspension feeders. Small fish landing on the upper surface may be captured by pedicilaria and dead animal matter may be scavenged but the main prey items are living invertebrates, mostly bivalve molluscs. To feed on one of these, the starfish moves over it, attaches its tube feet and exerts pressure on the valves by arching its back. When a small gap between the valves is formed, the starfish inserts part of its stomach into the prey, excretes digestive enzymes and slowly liquefies the soft body parts. As the adductor muscle of the shellfish relaxes, more stomach is inserted and when digestion is complete, the stomach is returned to its usual position in the starfish with its now liquefied bivalve meal inside it. The same everted stomach process is used by other starfish to feed on sponges, sea anemones, corals, detritus and algal films. [82]

Defense mechanisms

Despite their low nutrition value and the abundance of indigestible calcite, echinoderms are the prey of many organisms, such as crabs, sharks, sea birds and other echinoderms. Defensive strategies employed include the presence of spines, toxins, which can be inherent or delivered through the tube feet, and the discharge of sticky entangling threads by sea cucumbers. Although most echinoderm spines are blunt, those of the crown-of-thorns starfish are long and sharp and can cause a painful puncture wound as the epithelium covering them contains a toxin. [6] Because of their catch connective tissue, which can change rapidly from a flaccid to a rigid state, echinoderms are very difficult to dislodge from crevices. Certain sea cucumbers have a cluster of cuvierian tubules which can be ejected as long sticky threads from their anus and entangle and permanently disable an attacker. Another defensive strategy sometimes adopted by sea cucumbers is to rupture the body wall and discharge the gut and internal organs. The animal has a great regenerative capacity and will regrow the lost parts later. [83] Starfish and brittle stars may undergo autotomy when attacked, an arm becoming detached which may distract the predator for long enough for the animal to escape. Some starfish species can "swim" away from what may be danger, foregoing the regrowth by not losing limbs. [84] It is not unusual to find starfish with arms of different sizes in various stages of regrowth. [85]

Ecology

Echinoderms are numerous and relatively large invertebrates and play an important role in marine, benthic ecosystems. [10] The grazing of sea urchins reduces the rate of colonization of bare rock by settling organisms but also keeps algae in check, thereby enhancing the biodiversity of coral reefs. The burrowing of sand dollars, sea cucumbers and some starfish stirs up the sediment and depletes the sea floor of nutrients. Their digging activities increases the depth to which oxygen can seep and allows a more complex ecological tier-system to develop. Starfish and brittle stars prevent the growth of algal mats on coral reefs, which might otherwise obstruct the filter-feeding constituent organisms. [86] Some sea urchins can bore into solid rock and this bioerosion can destabilise rock faces and release nutrients into the ocean. Coral reefs are also bored into in this way but the rate of accretion of carbonate material is often greater than the erosion produced by the sea urchin. [87] It has been estimated that echinoderms capture and sequester about 0.1 gigatonnes of carbon per year as calcium carbonate, making them important contributors in the global carbon cycle. [88]

Echinoderms sometimes have large population swings which can cause marked consequences for ecosystems. An example is the change from a coral-dominated reef system to an alga-dominated one that resulted from the mass mortality of the tropical sea urchin Diadema antillarum in the Caribbean in 1983. [89] Sea urchins are among the main herbivores on reefs and there is usually a fine balance between the urchins and the kelp and other algae on which they graze. A diminution of the numbers of predators (otters, lobsters and fish) can result in an increase in urchin numbers causing overgrazing of kelp forests with the result that an alga-denuded "urchin barren" forms. [90] On the Great Barrier Reef, an unexplained increase in the numbers of crown-of-thorns starfish (Acanthaster planci), which graze on living coral tissue, has had considerable impact on coral mortality and coral reef biodiversity. [91]

Echinoderms form part of the diet of many organisms such as bony fish, sharks, eider ducks, gulls, crabs, gastropod molluscs, sea otters, Arctic foxes and humans. Larger starfish prey on smaller ones and the great quantity of eggs and larvae produced form part of the zooplankton, consumed by many marine creatures. Crinoids are relatively free from predation. [86] The body cavities of many sea cucumbers and some starfish provide a habitat for parasitic or symbiotic organisms including fish, crabs, worms and snails. [92]

Use by humans

In 2010, 373,000 tonnes of echinoderms were harvested, mainly for consumption. These were mainly sea cucumbers (158,000 tonnes) and sea urchins (73,000 tonnes). [93] [ clarification needed ]

Sea cucumbers are considered a delicacy in some countries of south east Asia; as such, they are in imminent danger of being over-harvested. [94]

Popular species include the pineapple roller Thelenota ananas (susuhan) and the red Holothuria edulis . These and other species are colloquially known as bêche de mer or trepang in China and Indonesia. The sea cucumbers are boiled for twenty minutes and then dried both naturally and later over a fire which gives them a smoky tang. In China they are used as a basis for gelatinous soups and stews. [95] Both male and female gonads of sea urchins are also consumed particularly in Japan, Peru, Spain and France. The taste is described as soft and melting, like a mixture of seafood and fruit. The quality is assessed by the colour which can range from light yellow to bright orange. [96] At the present time, some trials of breeding sea uchins in order to try to compensate the overexploitation of this resource have been made. [97]

The calcareous tests or shells of echinoderms are used as a source of lime by farmers in areas where limestone is unavailable and some are used in the manufacture of fish meal. [98] Four thousand tons of the animals are used annually for these purposes. This trade is often carried out in conjunction with shellfish farmers, for whom the starfish pose a major threat by eating their cultured stock. Other uses for the starfish they recover include the manufacture of animal feed, composting and drying for the arts and craft trade. [99]

Sea urchins are used in research, particularly as model organisms in developmental biology [100] and ecotoxicology. [101] [102] Strongylocentrotus purpuratus and Arbacia punctulata are used for this purpose in embryological studies. [103] The large size and the transparency of the eggs enables the observation of sperm cells in the process of fertilising ova. [100] The arm regeneration potential of brittle stars is being studied in connection with understanding and treating neurodegenerative diseases in humans. [99]

See also

Related Research Articles

Crinoid Class of echinoderms

Crinoids are marine animals that make up the class Crinoidea, one of the classes of the phylum Echinodermata, which also includes the starfish, brittle stars, sea urchins and sea cucumbers. Those crinoids which, in their adult form, are attached to the sea bottom by a stalk are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, being members of the largest crinoid order, Comatulida.

Sea urchin Class of echinoderms

Sea urchins, or simply urchins, are typically spiny, globular animals, echinoderms in the class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal to 5,000 metres. Their tests are round and spiny, typically from 3 to 10 cm across. Sea urchins move slowly, crawling with their tube feet, and sometimes pushing themselves with their spines. They feed primarily on algae but also eat slow-moving or sessile animals. Their predators include sea otters, starfish, wolf eels, and triggerfish.

Sea cucumber class of echinoderms

Sea cucumbers are echinoderms from the class Holothuroidea. They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothurian species worldwide is about 1,717 with the greatest number being in the Asia Pacific region. Many of these are gathered for human consumption and some species are cultivated in aquaculture systems. The harvested product is variously referred to as trepang, namako, bêche-de-mer or balate. Sea cucumbers serve a useful role in the marine ecosystem as they help recycle nutrients, breaking down detritus and other organic matter after which bacteria can continue the degradation process.

The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms move by alternately contracting muscles that force water into the tube feet, causing them to extend and push against the ground, then relaxing to allow the feet to retract.

Pedicellaria

A pedicellaria is a small wrench- or claw-shaped appendage with movable jaws, called valves, commonly found on echinoderms, particularly in sea stars and sea urchins. Each pedicellaria is an effector organ with its own set of muscles, neuropils, and sensory receptors and is therefore capable of reflex responses to the environment. Pedicellariae are poorly understood but in some taxa, they are thought to keep the body surface clear of algae, encrusting organisms, and other debris in conjunction with the ciliated epidermis present in all echinoderms.

Brittle star class of echinoderms

Brittle stars or ophiuroids are echinoderms in the class Ophiuroidea closely related to starfish. They crawl across the sea floor using their flexible arms for locomotion. The ophiuroids generally have five long, slender, whip-like arms which may reach up to 60 cm (24 in) in length on the largest specimens. They are also known as serpent stars; the New Latin class name Ophiuroidea is derived from the Ancient Greek ὄφις, meaning "serpent".

Tube feet Multipurpose organs of echinoderms

Tube feet are small active tubular projections on the oral face of an echinoderm, whether the arms of a starfish, or the undersides of sea urchins, sand dollars and sea cucumbers; they are more discrete though present on brittlestars, and have only a feeding function in feather stars. They are part of the water vascular system.

<i>Synaptula lamperti</i> Species of echinoderm

Synaptula lamperti is a species of sea cucumber in the family Synaptidae in the phylum Echinodermata, found on coral reefs in the Indo-Pacific region. The echinoderms are marine invertebrates and include the sea urchins, starfish and sea cucumbers. They are radially symmetric and have a water vascular system that operates by hydrostatic pressure, enabling them to move around by use of many suckers known as tube feet. Sea cucumbers are usually leathery, gherkin-shaped animals with a cluster of short tentacles at one end. They live on the sea bottom.

<i>Luidia ciliaris</i> species of echinoderm

The seven-armed sea star is a species of sea star (starfish) in the family Luidiidae. It is found in the eastern Atlantic Ocean and the Mediterranean Sea.

<i>Metacrinus rotundus</i> species of echinoderm

Metacrinus rotundus, the Japanese sea lily, is a marine invertebrate, a species of stalked crinoid in the family Isselicrinidae. It is a species found off the west coast of Japan, and is living near the edge of the continental shelf, around 100-150m deep. This is the shallowest species among the extant stalked crinoids.

Holothuria parvula, the golden sea cucumber, is a species of echinoderm in the class Holothuroidea. It was first described by Emil Selenka in 1867 and has since been placed in the subgenus Platyperona, making its full scientific name Holothuria (Platyperona) parvula. It is found in shallow areas of the Caribbean Sea and Gulf of Mexico and is unusual among sea cucumbers in that it can reproduce by breaking in half.

Ossicle (echinoderm)

Ossicles are small calcareous elements embedded in the dermis of the body wall of echinoderms. They form part of the endoskeleton and provide rigidity and protection. They are found in different forms and arrangements in sea urchins, starfish, brittle stars, sea cucumbers, and crinoids. The ossicles and spines are the only parts of the animal likely to be fossilized after an echinoderm dies.

<i>Ophiactis savignyi</i> species of echinoderm

Ophiactis savignyi is a species of brittle star in the family Ophiactidae, commonly known as Savigny's brittle star or the little brittle star. It occurs in the tropical and subtropical parts of all the world's oceans and is thought to be the brittle star with the most widespread distribution. It was first described by the German zoologists Johannes Peter Müller and Franz Hermann Troschel in 1842. The specific name honours the French zoologist Marie Jules César Savigny.

<i>Neoferdina cumingi</i> species of echinoderm

Neoferdina cumingi, also known as Cuming's sea star, is a species of starfish in the family Goniasteridae. It is native to the tropical Indo-Pacific region.

<i>Coscinasterias muricata</i> species of echinoderm

Coscinasterias muricata is a species of starfish in the family Asteriidae. It is a large 11-armed starfish and occurs in shallow waters in the temperate western Indo-Pacific region.

<i>Aporometra wilsoni</i> species of echinoderm

Aporometra wilsoni is a marine invertebrate, a species of crinoid or feather star in the family Aporometridae. It is found in shallow water around the coasts of southern Australia.

References

  1. Stöhr, Sabine (2014). "Echinodermata". WoRMS. World Register of Marine Species . Retrieved 23 February 2014.
  2. 1 2 "echinoderm". Online Etymology Dictionary .
  3. "Sea Lily". Science Encyclopedia. Retrieved 5 September 2014.
  4. "Animal Diversity Web - Echinodermata". University of Michigan Museum of Zoology. Retrieved 26 August 2012.
  5. "Computer simulations reveal feeding in early animal".
  6. 1 2 Dorit, R. L.; Walker, W. F.; Barnes, R. D. (1991). Zoology. Saunders College Publishing. pp. 777–779. ISBN   978-0-03-030504-7.
  7. Richard Fox. "Asterias forbesi". Invertebrate Anatomy OnLine. Lander University. Retrieved 19 May 2012.
  8. 1 2 Wray, Gregory A. (1999). "Echinodermata: Spiny-skinned animals: sea urchins, starfish, and their allies". Tree of Life web project. Retrieved 19 October 2012.
  9. Telford, M. J.; Lowe, C. J.; Cameron, C. B.; Ortega-Martinez, O.; Aronowicz, J.; Oliveri, P.; Copley, R. R. (2014). "Phylogenomic analysis of echinoderm class relationships supports Asterozoa". Proceedings of the Royal Society B: Biological Sciences. 281 (1786): 20140479–20140479. doi:10.1098/rspb.2014.0479. PMC   4046411 .
  10. 1 2 3 4 Uthicke, Sven; Schaffelke, Britta; Byrne, Maria (1 January 2009). "A boom–bust phylum? Ecological and evolutionary consequences of density variations in echinoderms". Ecological Monographs. 79: 324. doi:10.1890/07-2136.1.
  11. Siera104. "Echinodermata" . Retrieved 15 March 2008.
  12. 1 2 3 Waggoner, Ben (16 January 1995). "Echinodermata: Fossil Record". Introduction to the Echinodermata. Museum of Paleontology: University of California at Berkeley. Retrieved 14 March 2013.
  13. Smith, Dave (28 September 2005). "Vendian Animals: Arkarua" . Retrieved 14 March 2013.
  14. 1 2 Dorit, Walker & Barnes (1991) p. 792–793
  15. UCMP Berkeley, edu. "Echinodermata: Morphology". University of California Museum of Paleontology. Retrieved 21 March 2011.
  16. 1 2 Ruppert, Fox & Barnes (2004) p. 873
  17. Australian Echinoderms: Biology, Ecology and Evolution
  18. Messing, Charles. "Crown and calyx". Charles Messing's Crinoid Pages. Retrieved 29 July 2012.
  19. Behrens, Peter; Bäuerlein, Edmund (2007). Handbook of Biomineralization: Biomimetic and bioinspired chemistry'. Wiley-VCH. p. 393. ISBN   3-527-31805-4.
  20. Davies, A. Morley (1925). An Introduction to Palaeontology. Thomas Murby. pp. 240–241.
  21. Weber, W.; Dambach, M. (April 1974). "Light-sensitivity of isolated pigment cells of the sea urchin Centrostephanus longispinus". Cell and Tissue Research. 148 (3): 437–440. doi:10.1007/BF00224270. PMID   4831958.
  22. Motokawa, Tatsuo (May 1984). "Connective tissue catch in echinoderms". Biological Reviews. 59 (2): 255–270. doi:10.1111/j.1469-185X.1984.tb00409.x.
  23. 1 2 Dorit, Walker & Barnes (1991) pp. 780–791
  24. Dorit, Walker & Barnes (1991) p. 784–785
  25. Dorit, Walker & Barnes (1991) p. 788–789
  26. Dorit, Walker & Barnes (1991) p. 780–783
  27. Ruppert, Fox & Barnes (2004) p. 905
  28. Echinoderm Nutrition
  29. Rigby, P. Robin; Iken, Katrin; Shirayama, Yoshihisa (1 December 2007). Sampling Biodiversity in Coastal Communities: NaGISA Protocols for Seagrass and Macroalgal Habitats. NUS Press. p. 44. ISBN   9789971693688.
  30. Ruppert, Fox & Barnes (2004) p. 885
  31. Ruppert, Fox & Barnes (2004) p. 891
  32. Ruppert, Fox & Barnes (2004) pp. 902–904
  33. Ruppert, Fox & Barnes (2004) p. 912
  34. Ruppert, Fox & Barnes (2004) p. 920
  35. J. Moore. An Introduction to the Invertebrates, Cambridge Univ. Press, 2nd ed., 2006, p. 245.
  36. C. Hickman Jr., L. Roberts, A. Larson. Animal Diversity. Granite Hill Publishers, 2003. p. 271.
  37. "Macrobenthos of the North Sea - Echinodermata > Introduction". etibioinformatics.nl.
  38. Nielsen, Claus. Animal Evolution: Interrelationships of the Living Phyla. 3rd ed. Oxford, UK: Oxford University Press, 2012, p. 78
  39. 1 2 Ramirez-Gomez, Fransisco (27 September 2010). "Echinoderm Immunity". Invertebrate Survival Journal.
  40. Smith, Courtney (January 2010). "Echinoderm Immunity". Advances in Experimental Medicine and Biology: 260–301. doi:10.1007/978-1-4419-8059-5_14.
  41. 1 2 Ruppert, Fox & Barnes (2004) pp. 872–929
  42. Edmondson, C.H (1935). "Autotomy and regeneration of Hawaiian starfishes" (PDF). Bishop Museum Occasional Papers. 11 (8): 3–20.
  43. 1 2 3 4 McAlary, Florence A (1993). Population Structure and Reproduction of the Fissiparous Seastar, Linckia columbiae Gray, on Santa Catalina Island, California. 3rd California Islands Symposium. National Park Service. Retrieved 15 April 2012.
  44. 1 2 3 4 See last paragraph in review above AnalysisHotchkiss, Frederick H. C. (1 June 2000). "On the Number of Rays in Starfish". American Zoologist. 40 (3): 340–354. doi:10.1093/icb/40.3.340 . Retrieved 14 July 2011.
  45. 1 2 3 4 Fisher, W. K. (1 March 1925). "Asexual Reproduction in the Starfish, Sclerasterias" (PDF). Biological Bulletin. 48 (3): 171–175. doi:10.2307/1536659. ISSN   0006-3185. JSTOR   1536659 . Retrieved 15 July 2011.
  46. Dobson, W. E.; S. E. Stancyk; L. A. Clements; R. M. Showman (1 February 1991). "Nutrient Translocation during Early Disc Regeneration in the Brittlestar Microphiopholis gracillima (Stimpson) (Echinodermata: Ophiuroidea)". Biol Bull. 180 (1): 167–184. doi:10.2307/1542439. JSTOR   1542439 . Retrieved 14 July 2011.
  47. Mashanov, Vladimir S.; Igor Yu. Dolmatov; Thomas Heinzeller (1 December 2005). "Transdifferentiation in Holothurian Gut Regeneration". Biol Bull. 209 (3): 184–193. doi:10.2307/3593108. JSTOR   3593108. PMID   16382166 . Retrieved 15 July 2011.
  48. Hart, M. W. (January–February 2002). "Life history evolution and comparative developmental biology of echinoderms". Evolutionary Development. 4 (1): 62–71. doi:10.1046/j.1525-142x.2002.01052.x. PMID   11868659.
  49. Young, Craig M.; Eckelbarger, Kevin J. (1994). Reproduction, Larval Biology, and Recruitment of the Deep-Sea Benthos. Columbia University Press. pp. 179–194. ISBN   0231080042.
  50. Ruppert, Fox & Barnes (2004) pp. 887–888
  51. Ruppert, Fox & Barnes (2004) p. 895
  52. Ruppert, Fox & Barnes (2004) p. 888
  53. Ruppert, Fox & Barnes (2004) p. 908
  54. Ruppert, Fox & Barnes (2004) p. 916
  55. Ruppert, Fox & Barnes (2004) p. 922
  56. Yamaguchi, M.; J. S. Lucas (1984). "Natural parthenogenesis, larval and juvenile development, and geographical distribution of the coral reef asteroid Ophidiaster granifer". Marine Biology. 83 (1): 33–42. doi:10.1007/BF00393083. ISSN   0025-3162.
  57. McGovern, Tamara M. (5 April 2002). "Patterns of sexual and asexual reproduction in the brittle star Ophiactis savignyi in the Florida Keys" (PDF). Marine Ecology Progress Series. 230: 119–126. doi:10.3354/meps230119.
  58. 1 2 3 Monks, Sarah P. (1 April 1904). "Variability and Autotomy of Phataria". Proceedings of the Academy of Natural Sciences of Philadelphia. 56 (2): 596–600. ISSN   0097-3157. JSTOR   4063000.
  59. Kille, Frank R. (1942). "Regeneration of the Reproductive System Following Binary Fission in the Sea-Cucumber, Holothuria parvula (Selenka)" (PDF). Biological Bulletin. 83 (1): 55–66. doi:10.2307/1538013. ISSN   0006-3185. JSTOR   1538013 . Retrieved 15 July 2011.
  60. 1 2 Eaves, Alexandra A.; Palmer, A. Richard (11 September 2003). "Reproduction: Widespread cloning in echinoderm larvae". Nature. 425 (6954): 146. doi:10.1038/425146a. ISSN   0028-0836. PMID   12968170.
  61. Jaeckle, William B. (1 February 1994). "Multiple Modes of Asexual Reproduction by Tropical and Subtropical Sea Star Larvae: An Unusual Adaptation for Genet Dispersal and Survival" (PDF). Biological Bulletin. 186 (1): 62–71. doi:10.2307/1542036. ISSN   0006-3185. JSTOR   1542036 . Retrieved 13 July 2011.
  62. 1 2 3 Vaughn, Dawn (October 2009). "Predator-Induced Larval Cloning in the Sand Dollar Dendraster excentricus: Might Mothers Matter?". Biological Bulletin. 217 (2): 103–114. doi:10.1086/BBLv217n2p103. PMID   19875816.
  63. McDonald, Kathryn A.; Dawn Vaughn (1 August 2010). "Abrupt Change in Food Environment Induces Cloning in Plutei of Dendraster excentricus". Biological Bulletin. 219 (1): 38–49. doi:10.1086/BBLv219n1p38. PMID   20813988 . Retrieved 16 July 2011.
  64. 1 2 Vaughn, Dawn; Richard R. Strathmann (14 March 2008). "Predators induce cloning in echinoderm larvae". Science. 319 (5869): 1503. doi:10.1126/science.1151995. JSTOR   40284699. PMID   18339931 . Retrieved 16 July 2011.
  65. Vaughn, Dawn (March 2010). "Why run and hide when you can divide? Evidence for larval cloning and reduced larval size as an adaptive inducible defense". Marine Biology. 157 (6): 1301–1312. doi:10.1007/s00227-010-1410-z. ISSN   0025-3162.
  66. 1 2 Dorit, R. L.; Walker, W. F.; Barnes, R. D. (1991). Zoology. Saunders College Publishing. p. 778. ISBN   978-0-03-030504-7.
  67. Wim van Egmond (1 July 2000). "Gallery of Echinoderm Larvae". Microscopy UK. Retrieved 2 February 2013.
  68. Xiaofeng, Wang; Hagdorn, Hans; Chuanshang, Wang (September 2006). "Pseudoplanktonic lifestyle of the Triassic crinoid Traumatocrinus from Southwest China". Lethaia. 39 (3): 187–193. doi:10.1080/00241160600715321.
  69. Pawson, David Leo. "Echinoderm: distribution and abundance". Encyclopædia Britannica. Retrieved 28 June 2013.
  70. 1 2 Smith, J. E. (1937). "The structure and function of the tube feet in certain echinoderms" (PDF). Journal of the Marine Biological Association of the United Kingdom. 22 (1): 345–357. doi:10.1017/S0025315400012042. Archived from the original (PDF) on 15 November 2013.Cite uses deprecated parameter |deadurl= (help)
  71. "Leather star - Dermasterias imbricata". Sea Stars of the Pacific Northwest. Archived from the original on 9 September 2012. Retrieved 27 September 2012.Cite uses deprecated parameter |dead-url= (help)
  72. "Sand star - Luidia foliolata". Sea Stars of the Pacific Northwest. Archived from the original on 9 September 2012. Retrieved 26 September 2012.Cite uses deprecated parameter |dead-url= (help)
  73. Ruppert, Fox & Barnes (2004) pp. 899–900
  74. Ruppert, Fox & Barnes (2004) pp. 911–912
  75. Messing, Charles. "The crinoid feeding mechanism". Charles Messing's Crinoid Pages. Retrieved 26 July 2012.
  76. Barnes, Robert D. (1982). Invertebrate Zoology. Holt-Saunders International. pp. 997–1007. ISBN   0-03-056747-5.
  77. Tom Carefoot. "Learn about feather stars: feeding & growth". A Snail's Odyssey. Retrieved 23 February 2013.
  78. Ruppert, Fox & Barnes (2004) p. 893
  79. Tom Carefoot. "Learn about sea urchins: feeding, nutrition & growth". A Snail's Odyssey. Retrieved 23 February 2013.
  80. Tom Carefoot. "Learn about sand dollars: feeding & growth". A Snail's Odyssey. Retrieved 23 February 2013.
  81. Ruppert, Fox & Barnes (2004) p. 914
  82. Ruppert, Fox & Barnes (2004) pp. 884–885
  83. Dorit, Walker & Barnes (1991) p. 789–790
  84. Mladenov, Philip V.; Igdoura, Suleiman; Asotra, Satish; Burke, Robert D. (1 April 1989). "Purification and partial characterization of an autotomy-promoting factor from the sea star Pycnopodia Helianthoides". The Biological Bulletin. 176 (2): 169–175. doi:10.2307/1541585. ISSN   0006-3185.
  85. Beamer, Victoria P. "Regeneration in Starfish (Asteroidea)". The Physiology of Arm Regeneration in Starfish (Asteroidea). Davidson College. Archived from the original on 9 April 2013. Retrieved 10 March 2013.Cite uses deprecated parameter |dead-url= (help)
  86. 1 2 Miller, John E. "Echinoderm: role in nature". Encyclopædia Britannica. Retrieved 28 June 2013.
  87. Herrera-Escalante, T; López-Pérez, R. A.; Leyte-Morales, G. E. (2005). "Bioerosion caused by the sea urchin Diadema mexicanum (Echinodermata: Echinoidea) at Bahías de Huatulco, Western Mexico". Revista de Biologia Tropical. 53 (3): 263–273. PMID   17469255.
  88. Kaplan, M. (2010). "Sea stars suck up carbon". Nature. doi:10.1038/news.2009.1041.
  89. Osborne, Patrick L. (2000). Tropical Ecosystem and Ecological Concepts. Cambridge: Cambridge University Press. p. 464. ISBN   0-521-64523-9.
  90. Lawrence, J. M. (1975). "On the relationships between marine plants and sea urchins" (PDF). Oceanographic Marine Biological Annual Review. 13: 213–286. Archived from the original (PDF) on 5 February 2011.Cite uses deprecated parameter |deadurl= (help)
  91. Birkeland, Charles; Lucas, John S. (1990). Acanthaster planci: Major Management Problem of Coral Reefs. Florida: CRC Press. ISBN   0849365996.
  92. Luciano, Brooke; Lyman, Ashleigh; McMillan, Selena; Nickels, Abby. "The symbiotic relationship between Sea cucumbers (Holothuriidae) and Pearlfish (Carapidae)" . Retrieved 27 June 2013.
  93. Sourced from the data reported in the FAO FishStat database
  94. "Sea Cucumbers Threatened by Asian Trade". New York Times. Retrieved 1 April 2009.
  95. Wikisource:1911 Encyclopædia Britannica/Bêche-de-Mer
  96. John M. Lawrence (2001). "The edible sea urchins". In John M. Lawrence (ed.). Edible Sea Urchins: Biology and Ecology. Developments in Aquaculture and Fisheries Science. 37. pp. 1–4. doi:10.1016/S0167-9309(01)80002-8. ISBN   9780444503909.
  97. Sartori D., Scuderi A.; Sansone G.; Gaion A. (February 2015). "Echinoculture: the rearing of Paracentrotus lividus in a recirculating aquaculture system—experiments of artificial diets for the maintenance of sexual maturation". Aquaculture International. 23: 111–125. doi:10.1007/s10499-014-9802-6.
  98. "Sea stars". Wild Singapore. Retrieved 4 February 2013.
  99. 1 2 Barkhouse, C.; Niles, M.; Davidson, L. -A. (2007). "A literature review of sea star control methods for bottom and off bottom shellfish cultures" (PDF). Canadian Industry Report of Fisheries and Aquatic Sciences. 279 (7): 14–15. ISSN   1488-5409.
  100. 1 2 "Insight from the Sea Urchin". Microscope Imaging Station. Exploratorium. Retrieved 4 February 2013.
  101. Sartori D., Gaion A. (2015). "Toxicity of polyunsaturated aldehydes of diatoms to Indo-Pacific bioindicator organism Echinometra mathaei". Drug and Chemical Toxicology: 1–5. doi:10.3109/01480545.2015.1041602.
  102. Gaion A., Scuderi A.; Pellegrini D.; Sartori D. (2013). "Arsenic Exposure Affects Embryo Development of Sea Urchin, Paracentrotus lividus (Lamarck, 1816)". Bulletin of Environmental Contamination and Toxicology: 1–5. doi:10.3109/01480545.2015.1041602.
  103. Longo, F. J.; Anderson, E. (June 1969). "Sperm differentiation in the sea urchins Arbacia punctulata and Strongylocentrotus purpuratus". Journal of Ultrastructure Research. 27 (5): 486–509. doi:10.1016/S0022-5320(69)80046-8. PMID   5816822.

Cited texts

Further reading