Introduction to the Echinodermata . . . from starfish to sea cucumbers. . . A nyone who has been to the beach has probably seen starfish or sand dollars. The more intrepid beachcomber may find brittle stars, sea cucumbers, or sea urchins. These and many other organisms, living and extinct, make up the Echinodermata, the largest phylum to lack any freshwater or land representatives. Most living echinoderms, like this sand dollar from Baja California, are pentameral; that is, they have fivefold symmetry, with rays or arms in fives or multiples of five. However, a number of fossil echinoderms were not pentameral at all, and some had downright bizarre shapes. Echinoderms have a system of internal water-filled canals, which in many echinoderms form suckered "tube feet", with which the animal may move or grip objects. Echinodermata By Renee Sherman Mulcrone Kingdom: Animalia : Echinodermata Members of this Diversity Echinodermata has approximately 7000 described living species and about 13,000 extinct species known from the fossil record. This phylum is the largest without any freshwater or terrestrial forms. (Brusca and Brusca, 2003; Waggoner, 1999) Geographic Range Mainly a marine group, echinoderms are found in all the oceans. (Brusca and Brusca, 2003) Biogeographic Regions arctic ocean /site/glossary/popup/20020917113601.html/site/glossary/popup/20020917113601.html; indian ocean; atlantic ocean /site/glossary/popup/20020917114102.html/site/glossary/popup/20020917114102.html; pacific ocean /site/glossary/popup/20020917114731.html/site/glossary/popup/20020917114731.html; mediterranean sea Habitat Except for a few species which inhabit brackish waters, all echinoderms are benthic organisms found in marine environments. Echinoderms inhabit depths ranging from shallow waters at tide lines to the deep sea. (Barnes, 1987; Brusca and Brusca, 2003; University of Alabama Center for Communication and Educational Technology, 2000; Waggoner, 1999) Habitat Regions temperate /site/glossary/popup/20020904145595.html/site/glossary/popup/20020904145595.html; tropical /site/glossary/popup/20020904145598.html/site/glossary/popup/20020904145598.html; polar /site/glossary/popup/20020904145371.html/site/glossary/popup/20020904145371.html; saltwater or marine /site/glossary/popup/20020904145582.html/site/glossary/popup/20020904145582.html Aquatic Biomes brackish water /site/glossary/popup/20020904145435.html/site/glossary/popup/20020904145435.html Other Habitat Features intertidal or littoral /site/glossary/popup/20020904145633.html/site/glossary/popup/20020904145633.html Physical Description Larvae range from a few millimeters to a few decimeters, while adults can range from less than 1 cm to 2 m. While adult forms are radially symmetrical, larval forms are always bilateral. The radial symmetry is secondarily derived. The pentaradial form, whether it has arms or not, has a central disc. An internal skeleton is present throughout members of the phylum. Ossicles, which make up the skeleton, are below an outer dermal layer. The skeletal and muscular arrangement varies among groups. Pedicellariae produced by the skeleton, are pincer-like structures. Found mainly in echinoids and asteroids, their function is debatable. They may be used to capture prey, clean, or hold items to disguise from predators. Echinoderms have a water vascular system consisting of a network of radial canals, which extend through each of the five extensions (arms or rays) of the animal. Each canal has a lateral connection which leads to a tube foot, which may be composed of three parts. Internally is the ampulla and externally is the podia. At the end of the podia is usually a sucker. Grooves with rows of podia extending from the mouth are called the ambiculacra . Between each ambiculacra is the interambulacrum. For groups of animals with "arms" (sea stars, for example), the interambulacrum is just the space between the ambiculacra. For other animals without furrows (sea cucumbers, for example), the areas are like the ambiculacra, but usually lack holes for the tube feet. The water vasuclar system opening, called a madreporite, lies on a particular interambulacrum. Letters are used to describe parts of echinoderms. The ambulacrum opposite the madreporite is section A. Moving clockwise, other parts are coded B through E. Sections C and D are termed the bivium while all the others are collectively termed the trivium. Interambulacrum sections are named using the letters of the ambulacra sections they are between (e. g. AB). (Barnes, 1987; Brusca and Brusca, 2003; Waggoner, 1999) Other Physical Features ectothermic /site/glossary/popup/20020916131519.html/site/glossary/popup/20020916131519.html; heterothermic /site/glossary/popup/20020916125902.html/site/glossary/popup/20020916125902.html; bilateral symmetry /site/glossary/popup/20020904145642.html/site/glossary/popup/20020904145642.html; radial symmetry /site/glossary/popup/20020904145808.html/site/glossary/popup/20020904145808.html Development Echinoderms are deuterostomes. The larvae, which are planktotrophic or lecithotrophic, have 3part paired coeloms. Embryonic coelomic structures have specific fates as the bilaterally symmetrical larvae metamorphose into radially symmetric adults. Adult pheromones may attract larvae, which tend to settle near conspecific adults. Metamorphosis in some species is triggered by adult pheromones. (Barnes, 1987; Brusca and Brusca, 2003) Development - Life Cycle metamorphosis /site/glossary/popup/20020904145364.html/site/glossary/popup/20020904145364.html Reproduction Mating System polygynandrous (promiscuous) /site/glossary/popup/20020904145483.html/site/glossary/popup/20020904145483.html Echinoderms are mainly gonochoristic (having separate sexes), with exceptions among the asteroids, holothurians and ophuroids. Holothurians possess a single gonad, crinoids lack distinct gonads, while asteroids and echinoids have multiple gonads. Echinoderm reproductive strategies vary from free spawning and indirect development to brooding and direct development. Spawning is probably a noctural event. (Barnes, 1987; Brusca and Brusca, 2003) Key Reproductive Features seasonal breeding /site/glossary/popup/20020904145584.html/site/glossary/popup/20020904145584.html; gonochoric/gonochoristic/dioecious (sexes separate); simultaneous hermaphrodite; sequential hermaphrodite; sexual /site/glossary/popup/20020904145786.html/site/glossary/popup/20020904145786.html; fertilization /site/glossary/popup/20020904145546.html/site/glossary/popup/20020904145546.html(External /site/glossary/popup/20020904145726.html/site/glossary/popup/20020904145726.html); oviparous /site/glossary/popup/20020904145572.html/site/glossary/popup/20020904145572.html Parental investment ranges from no care after the release of eggs for free spawning to brooding the young. Brooding is found in polar and boreal echinoderms and some deep sea echinoderms, where environments are more difficult for the larvae. (Barnes, 1987; Brusca and Brusca, 2003) Parental Investment pre-fertilization (Provisioning); pre-hatching/birth (Provisioning: Female, Protecting: Female) Behavior Most radially symmetric animals are sessile, however, echinoderms are able to move. The water vascular system originally functioned for collection and transport of food, but evolved to function for locomotion as well. (Barnes, 1987; Brusca and Brusca, 2003) Key Behaviors diurnal /site/glossary/popup/20020904145414.html/site/glossary/popup/20020904145414.html; nocturnal /site/glossary/popup/20020904145503.html/site/glossary/popup/20020904145503.html; motile /site/glossary/popup/20020904145472.html/site/glossary/popup/20020904145472.html; sedentary /site/glossary/popup/20020904145585.html/site/glossary/popup/20020904145585.html Communication and Perception The non-centralized nervous system allows echinoderms to sense their environment from all sides. Adult pheromones may attract larvae, which tend to settle near conspecific adults. Metamorphosis in some species is triggered by adult pheromones. (Brusca and Brusca, 2003) Communication Channels chemical /site/glossary/popup/20020904145606.html/site/glossary/popup/20020904145606.html Other Communication Modes pheromones /site/glossary/popup/20020904145769.html/site/glossary/popup/20020904145769.html Perception Channels tactile /site/glossary/popup/20020904145500.html/site/glossary/popup/20020904145500.html; chemical /site/glossary/popup/20020904145606.html/site/glossary/popup/20020904145606.html Food Habits Different groups have different feeding habits. Members of the Crinoidea sit with arms outstretched parallel to the currents and filter feed on passing particles. Most Asteroidea are predators or scavengers, everting their stomach (called a cardiac stomach), which secretes digestive enzymes on their prey. Some asteroids are also suspension feeders. Brittle stars of the Ophiuroidea are predators, deposit feeders, scavengers, and suspension feeders, which feed by outstretching their arms to capture prey. Ophiuroids lack an intestine and anus, and therefore have an incomplete digestive system. The members of Echinoidea are suspension feeders, herbivores, detritivores, and predators. Many have a group of hard plates which retract and grasp like teeth, commonly called Aristotle's lantern. This allows most sea urchins to graze on algae. Most Holothuroidea are suspension or deposit feeders. Holothurians may also eviserate their digestive and other organs in response to predation or seasonal events. (Brusca and Brusca, 2003; Waggoner, 1999) Foraging Behavior filter-feeding /site/glossary/popup/20020904145660.html/site/glossary/popup/20020904145660.html Predation Known Predators otters (Lutrinae) flounders (Pleuronectidae) haddock (Melanogrammus) Echinoderms in general are most vulnerable in their larval stage. As adults, asteroids have an anti-predator adaptation where they can lose an arm to a predator and the arm is later regenerated. Holothurians discharge sticky tubules, known as Cuvierian tubules , at a potential predator. Otters prey mainly on sea urchins. (Brusca and Brusca, 2003) Ecosystem Roles Echinoderms are usually intricate parts of their ecosystems. Many asteroids are keystone species. Sea urchins, if not controlled by predators, may overgraze their habitat. Asteroids have several commensals, including polychaetes that feed on leftovers from the sea star's prey items. (Barnes, 1987; Brusca and Brusca, 2003) Ecosystem Impact keystone species /site/glossary/popup/20020904145561.html/site/glossary/popup/20020904145561.html Economic Importance for Humans: Positive Research on echinoderms has contributed to the overall knowledge of animal fertilization and development. Many echinoderms are easy to culture and maintain in a lab setting, and produce a large amount of eggs. Sea urchin eggs are also edible and often served in sushi bars. (Brusca and Brusca, 2003; University of Alabama Center for Communication and Educational Technology, 2000) Positive Impacts food /site/glossary/popup/20020904145418.html/site/glossary/popup/20020904145418.html; research and education Conservation Status The European edible sea urchin, Echinus esculentus, is listed as endangered by the IUCN. Isostichopus fuscus, a holothurian, is listed by CITES. It occurs on the coasts of Ecuador, Galapagos, Mexico and Peru. (UNEP-WCMC, 2005; World Conservation Monitoring Centre, 2004) Other Comments Since they are almost exclusively marine species, echinoderms are probably osmoconformers, with little ionic regulation. (Brusca and Brusca, 2003) For More Information Find Echinodermata information at Encyclopedia of Life Contributors Renee Sherman Mulcrone (author), . References Barnes, R. 1987. Invertebrate Zoology. Orlando, Florida: Dryden Press. Brusca, R., G. Brusca. 2003. Invertebrates. Sunderland, Massachusetts: Sinauer Associates, Inc.. UNEP-WCMC, 2005. "Isostichopus fuscus" (On-line). UNEP-WCMC Species Database: CITES-Listed Species. Accessed January 21, 2005 at http://www.cites.org/eng/resources/species.html. University of Alabama Center for Communication and Educational Technology, 2000. "Phylum Echinodermata – echinoderms" (On-line). Accessed January 16, 2005 at http://www.ccet.ua.edu/expedition/scsstarsurcbrit.htm. University of Paisley, 1998. "Echinodermata" (On-line). Accessed January 16, 2005 at http://www-biol.paisley.ac.uk/courses/Tatner/biomedia/units/echi1.htm. Waggoner, B. 1999. "Introduction to the Echinodermata" (On-line). Accessed January 16, 2005 at http://www.ucmp.berkeley.edu/echinodermata/echinodermata.html. World Conservation Monitoring Centre, 2004. "Echinus esculentus" (On-line). 2004 IUCN Red List of Threatened Species. Accessed January 21, 2005 at http://www.iucnredlist.org/search/details.php?species=7011. Chordates are defined as organisms that possess a structure called a notochord, at least during some part of their development. The notochord is a rod that extends most of the length of the body when it is fully developed. Lying dorsal to the gut but ventral to the central nervous system, it stiffens the body and acts as support during locomotion. Other characteristics shared by chordates include the following (from Hickman and Roberts, 1994): bilateral symmetry segmented body, including segmented muscles three germ layers and a well-developed coelom. single, dorsal, hollow nerve cord, usually with an enlarged anterior end (brain) tail projecting beyond (posterior to) the anus at some stage of development pharyngeal pouches present at some stage of development ventral heart, with dorsal and ventral blood vessels and a closed blood system complete digestive system bony or cartilaginous endoskeleton usually present. Kingdom: Animalia Phylum: Chordata Members of this Phylum