Phylum Echinodermata The echinoderms (“hedgehog skin”) are a very unusual group that includes about 7000 living species. Members include: starfish, brittle stars, sea urchins, sea cucumbers, and sea lilies or feather stars. They are deuterostomes (as are chordates), but have secondarily evolved radial symmetry from bilateral symmetry (they still have bilaterally symmetrical larvae). Characteristics of the Echinodermata Exclusively a marine group. They cannot osmoregulate so rarely occur even in brackish water. The body is not segmented, but shows pentaradial symmetry. There is no head or brain and the nervous system is relatively simple. Characteristics of the Echinodermata They possess an endoskeleton of dermal calcareous ossicles, which are connected together by connective tissue. Possess a unique water vascular system that consists of a series of canals that extend from the body surface as tube feet. These tube feet are tentacle-like and enable the animal to move. In some species movement of the arms or spines contributes to locomotion too. Classes of Echinoderms There are a total of five classes of echinoderms and about 7300 species. Class Asteroidea: sea stars or starfishes Class Ophiuroidea: Brittle stars Class Echinoidea: Sea Urchins, Sand dollars Class Holothuroidea: Sea cucumbers Class Crinoidea: Sea Lilies and Feather stars. Class Asteroidea Star fish are the most familiar of the echinoderms and demonstrate the characteristics of the group well. Typically, they have 5 tapering arms, which merge gradually with the central disc. The mouth is found on the oral surface, the opposite side of the animal being referred to as the aboral surface. Cushion seastar (Asteroidea) Class Asteroidea The aboral surface of starfish is usually rough to the touch and has spines (although these may be flattened so the animal appears smooth.). Around the base of the spines are small pincerlike structures called pedicellariae, which are under muscular control and help to remove debris and protect the animals papulae (skin gills) projections of the coelomic cavity that increase the surface area for gas exchange. Figure 22.01 Figure 22.04f Pedicellariae Water vascular system Grooves called ambulacral grooves radiate out from the mouth on the oral surface. Tube feet project from the grooves. These tube feet are connected to the water vascular system, which is a unique invention of the echinoderms. Figure 22.02b Water vascular system The water vascular system is a compartment of the coelom and is a system of canals and tube feet. The water vascular system uses hydraulic pressure to extend, move and contract the tube feet enabling the starfish to move and feed. Water vascular system The water vascular system opens to the outside via small pores in a structure called the madreporite on the aboral surface. The madreporite connects to a ring canal and from this ring canal, radial canals extend down the abulacral groove of each arm. Figure 22.03b Water vascular system From each of the radial canals smaller lateral canals branch off, which connect to the tube feet. Each tube foot is a hollow, muscle-lined tube, which has a bulb at one end (on the inside of the arm) and a sucker at the other end which protrudes from the animal. Figure 22.03b Figure 22.03c Water vascular system The tube feet use hydraulics to work. Each tube foot contains a lot of connective tissue in its wall that maintains the tube at a fairly constant diameter. Contraction of muscles in the ampulla forces fluid into the tube foot which extends (a valve prevents fluid flowing back into the lateral canal). Water vascular system To withdraw the foot, muscles in the tube foot contract which pushes fluid back into the ampulla. The tube foot can bend to one side as a result of the contraction of muscles on one or other side of the tube foot. In addition, small muscles at the end of the tube foot can contract to raise the center of the tube foot and create a suction cup. Water vascular system As a result of the combination of hydraulics and muscular control the tube feet can be moved with a high degree of control and can also exert a strong pull. This enables the starfish to move up hard vertical surfaces and open up prey such as bivalves. On sediments suction doesn’t work well and under these conditions the tube feet are used more like legs. Feeding and digestion The mouth of a sea star opens into a twopart stomach in the central disk of the animal. The lower cardiac stomach can be everted and this ability is important in sea stars ability to consume certain prey. Feeding and digestion The cardiac stomach is connected to a pyloric stomach above it which in turn connects to digestive cecae in the arms where most extracellular digestion takes place. A short intestine leads from the pyloric stomach to the anus on the aboral side. Figure 22.03a Feeding and digestion Most sea stars are quite unselective carnivores and feed on molluscs, crustaceans, annelids, other invertebrates and other echinoderms. Some sea stars are major predators of economically important species of bivalve such as mussels and oysters. Feeding and digestion When a sea star attacks a bivalve it wraps itself around it and uses its tube feet to try to pull the two shells apart. The adductor muscles of the bivalve keep them closed, but the sea star can maintain its pull for long periods and eventually (after about half an hour) the adductor muscles of the bivalve become fatigued and the shells separate a little. Feeding and digestion Once a gap opens the sea star everts its soft flexible stomach into the gap and begins to secrete digestive enzymes that slowly break down the soft tissues of the bivalve Figure 22.05a Reproduction and development Most sea stars have separate sexes and fertilization is external. Eggs are brooded by some species, but in most cases the eggs are released to develop into larvae. Early development is typical of deuterostomes and the free swimming bipinnaria larva is bilaterally symetrical. Figure 22.09a Reproduction and development Late in development, however, the larva follows a path quite different from other deuterostomes. The bipinnaria larva transforms into a brachiolaria by growing three adhesive arms and a sucker at its anterior end. It then attaches to a substrate and undergoes metamorphosis. Figure 22.09b Reproduction and development During metamorphosis the larva is radically altered. The anterioposterior axis is lost and what was the left side becomes the oral surface and what was the right side becomes the aboral surface. Reproduction and development The existing mouth and anus disappear and new ones are produced in the appropriate locations. Finally short arms and podia develop and the animal detaches from its stalk to become a young starfish. Figure 22.08 Regeneration Echinoderms have the capacity to regenerate lost parts. For example, a sea star can regenerate a leg (or even all of its legs) if it is lost). If an arm that is removed contains at least one fifth of the central disk a new individual will develop from it. Class Ophiuroidea The Ophiuroidea includes the brittle stars and basket stars. This is the largest class of echinoderms with more than 2000 living species and it probably is also the most abundant group. They are very common in all kinds of benthic marine environments and abound on the abyssal sea floor in many places. Class Ophiuroidea Although similar to sea stars in having five arms, the brittle stars can be readily distinguished by their proportionally much longer and thinner arms and by the clear separation between the arms and the central disk. Brittle star (Ophiuroidea) Class Ophiuroidea Brittle stars move in a different manner to starfish. Their ambulacral grooves are covered over with interlocking ossicles. They possess tube feet which play a role in feeding, but don’t contribute to movement. Movement instead occurs by movement of the jointed arms. Class Ophiuroidea The mouth is surrounded by five moveable plates that act as jaws. Brittle stars lack intestines and an anus and waste exits via the mouth. Brittle stars feed on small particles that they glean from the sea bottom or particles they sieve out of the water with mucus strands attached to spines on their arms Figure 22.11 Class Echinoidea The Echinoidea include the familiar sea urchins, sand dollars, and heart urchins. They have a compact body that is enclosed in a calcareous shell (or test), which is formed from dermal ossicles that have become a series of 10 double rows of close-fitting plates. These plates also bear moveable, stiff, often long spines. Figure 22.15 Sea urchins 0126.jpg Sand dollar Figure 22.18a Class Echinoidea The sea urchins are considered to be “regular” echinoids because they are hemispherical and radially symmetrical. However, the sand dollars and heart urchins are considered “irregular” because they have become secondarily bilaterally symmetrical. Class Echinoidea In sand dollars the anus and mouth are located on the oral side with the mouth near the middle, but the anus shifted near the margin, so that an anterioposterior axis is recognizable. This is even more apparent in the heart urchins where the mouth is moved towards the anterior end and the anus (periproct) near the posterior end. Figure 22.17 Class Echinoidea Echinoids are widely distributed in all seas. Sea urchins are most typically found on rocky substrates, but sand dollars and heart urchins prefer sandy substrates. Sea urchins feed by grazing on algae growing on rocks and they do so using a complex structure called an Aristotle’s lantern. Aristotle’s Lantern The Aristotle’s lantern is a chewing mechanism that can be raised and lowered by muscles inside the urchin. The lantern consists of five plates connected to each other with connective tissue in a vase-like arrangement and each plate has a tooth on the end. Figure 22.19 Aristotle’s Lantern The lantern can be lowered to scrape off algae and then retracted inside using paired muscles. The Aristotle’s lantern connects directly to the esophagus, which connects in turn to the rest of the gut. Class Holothuroidea The Holothuroidea or sea cucumbers are a quite odd group whose members at first glance don’t seem that similar to the other echinoderms. Sea cucumbers are very elongated along the oral-aboral axis and have an array of oral tentacles (which are modified tube feet) around the mouth. Sea cucumber Class Holothuroidea Holothuroideans have a leathery body as the ossicles are small. Because they lie on one side, sea cucumbers typically have well developed tube feet on only the three ambulacra in contact with the substrate. The other two ambulacra have less well developed tube feet, which are not used for movement, but may play a sensory role. Class Holothuroidea Sea cucumbers are sluggish and don’t move quickly. They feed either on deposits, which they collect as they crawl along the sediment or are filter feeders that filter small particles from the water. Deposit feeding sea cucumbers are a successful group and make up as much as 90% of the biomass on the surface of the deep-sea. Class Holothuroidea An unusual feature of sea cucumbers is their respiratory tree, which is a many branched internal respiratory structure that connects to the cloaca. Water is pumped in and out of the tree by the muscular cloaca and the tree itself and gas exchange takes place. Figure 22.22 Class Holothuroidea Attached to the respiratory tree are structures called Cuverian tubules. These are used in defense and are discharged out of the animal if it is disturbed. The tubules are long, sticky and sometimes toxic and can entangle an attacker. Some species also may discharge their digestive tract, respiratory tree or gonads. These structures can be regenerated. Figure 22.23c Class Crinoidea Includes the sea lilies and feather stars, which have an extensive fossil record and once were much more abundant in the seas. The crinoids have a very feathery appearance that results from the branching of their five arms to produce many more arms each of which has many lateral pinnules. Figure 22.25 Class Crinoidea Crionoids occur most commonly in deep water, but there are shallow water species too. Not surprisingly with their appearance they filter feed and the food is carried down the ambulacral grooves to the mouth by cilia and tube feet. Figure 22.24 Class Crinoidea Crinoids have a similar water vascular system to other echinoderms and have only simple sense organs. They lack spines, a madreporite and pedicellariae. Most living species are not more than 12 inches long, but some fossil species had stalks over 75 feet long. Phylum Hemichordata The hemichordates are deuterostome marine animals that were previously classified as a subphylum of the chordates because they possess a dorsal nerve cord, gills slits and a rudimentary notochord. However, there is now consensus that the hemichordate “notochord” is not homologous with that of the chordates. Thus, they have been classified in their own phylum. Phylum Hemichordata The hemichordates are wormlike organisms that live on the bottom and are usually found in shallow waters. There are two classes: the Enteropneusta (the acorn worms) and the Pterobranchia. Acorn worms The acorn worms are deposit or suspension feeders that use their large proboscis to trap food in mucus that is then transported by cilia to the mouth. Figure 22.29a Saccoglossus an acorn worm (Hemichordata: class Enteropneusta) Acorn worms The larva of acorn worms is called a tornaria and closely resembles the bipinnaria larva of echinoderms, which supports their classification as close relatives of the echinoderms. Figure 22.31 (Hemichordate) (Echinoderm) Pterobranchia The pterobranchs are small colonial animals (17mm in length). They live in tubes which they produce and can freely move about in filter feeding using their arms, which are covered in tentacles. They lack a dorsal nerve cord, but one genus has a pair of gill slits. Figure 22.32 Cephalodiscus a pterobranch hemichordate Taxonomy of hemichordates Hemichordates were once classified among the chordates on the basis of their gill slits and dorsal nerve cord. More recently molecular sequence work that has examined Hox genes and rRNA places the henichordates in a clade with the echinoderms. This is supported by the similarities in larvae. Taxonomy of hemichordates In addition, research on an extinct goup called the carpoids has found gill slits which suggests that gill slits are ancestral for all deuterostomes. Figure 22.34