Cnidaria Cnidaria The phylum Cnidaria includes over 9,000 species of aquatic, radially symmetrical animals which have specialized stinging organelles called nematocysts. They include the jellyfish, box jellyfish, sea anemones, fire corals, sea pens and hard corals. Cnidaria Cnidarians are diploblastic having only two well-defined germ layers (ectoderm and endoderm). Cnidarians are the simplest animals equipped with nerve cells which are arranged into a nerve net, but there is no central nervous system. Nerve net The nerve net is a diffuse nervous system. Nerve impulses are transmitted between cells by the release of neurotransmitters from vesicles, which carry the signal across the synapse (gap) between cells. Unlike “higher animals” impulses can travel in both directions along a nerve because many synapses have vesicles on both sides. Nerve net In cnidarians there is no brain, but in some medusae there are multiple nerve nets. For example, in Scyphozoan jellyfish there is a fast-conducting nerve net for coordinating swimming movements and a slower net to coordinate movements of tentacles. Cnidaria: digestion Cnidarians have an internal body cavity, the gastrovascular cavity, but no one-way gut. Food enters and waste exits through the same opening, the oral cavity. Digestion takes place extracellularly within the gastrovascular cavity. Cnidaria: Body wall The body wall that surrounds the gastrovascular cavity has an outer epidermis and an inner gastrodermis. In between these two layers is a gelatinous layer of mesoglea, which may contain elastic fibers or be stiffened with spicules or flexible proteins. Figure 13.02 Fig. 7.2 Body forms Cnidaria have one of two basic body forms Polyp Medusa In some groups one or other body form is used exclusively, but in others the two forms are used in a single life cycle. Polyp and medusa The polyp or hydroid form is adapted to a sessile existence and the medusa form to a free-floating or pelagic life. In both cases radial symmetry is favored because stimuli and food are equally likely to come from all directions. Foraging strategy Cnidarians are sit-and-wait predators rather than pursuit predators. They wait for prey to drift or blunder into their tentacles, which in the case of jellyfish, are almost invisible. Polyp and medusa Polyps and medusae may look quite different, but are basically inverted versions of each other. Figure 13.02 Fig. 7.2 Polyp and medusa Both polyps and medusae are equipped with tentacles around the oral cavity. The tentacles are equipped with cnidocytes that contain stinging nematocysts, which are used to kill prey. Cnidocytes Cnidocytes contain cnidae which are stinging organelles (the most common of which is the nematocyst). The cnida is a small capsule made of chitin that contains a coiled, often barbed, filament. The capsule is covered by a lid, which has an associated trigger mechanism. Figure 13.03 Figure 7.3 Cnidocytes When the trigger mechanism is tripped the cnida is expelled at high velocity into the prey. In many cases a toxin is injected, but in others, the cnida entangles or sticks to the prey. Cnidocytes Nematocysts of most cnidarians are not harmful to humans, but the stings of some (e.g. Portugese Man-of-war) are painful or even fatal (certain box jellyfish). Figure 13.05 7.5 Movement of medusae The medusal form of cnidarians is pelagic and the medusa can move by rhythmically contracting and pulsing its bell, which expels water and drives the medusa upwards. Most cnidarians are relatively weak swimmers, but cubozoans (box jellyfish) swim strongly. Youtube videos Mudusae in aquaria http://www.youtube.com/watch?v=gSq72 gkTdH4&NR http://www.youtube.com/watch?v=0ANt1l LDtQw What do jellyfish eat? Jellyfish prey on anything small that their tentacles can kill. They eat the planktonic larvae of various organisms, crustaceans, small fish, fish eggs and other jellyfish. What eats jellyfish? Jellyfish are eaten by a variety of animals including other jellyfish, tuna, sharks, swordfish and turtles. Turtles, which face numerous other threats, frequently starve as a result of eating plastic bags that they mistake for jellyfish. What eats jellyfish? Populations of all predators of jellyfish have decreased dramatically in the past 100 years as a result of overfishing and mortality as a result of by-catch. Many shark populations have declined by >90% and some by as much as 99%. Many turtle populations have been pushed to the brink of extinction Jellyfish populations Jellyfish populations have increased dramatically in many oceans. Reduced predation has certainly played a role in that increase. Jellyfish populations It has been speculated that jellyfish numbers may have increased as a result of overfishing of various fish stocks. Because jellyfish consume the same food as many adult and larval fish they may be benefiting from the reduction in competition. Jellyfish populations Jellyfish populations also benefit from the fact that they are more tolerant of low oxygen conditions and high nutrient ocean conditions. Anoxic (no/low oxygen) conditions and dead zones as a result of fertilizer pollution and associated plankton blooms are common in the oceans today. Jellyfish populations As much as 1% of the oceans (especially in relatively enclosed bodies of water) are seasonal dead zones. These occur when high levels of fertilizer runoff lead to blooms of phytoplankton. When these die their decomposition removes oxygen from the water and this kills a wide variety of organisms. Jellyfish populations Jellyfish, however, can thrive low oxygen conditions. In addition in high nutrient conditions with lots of phytoplankton smaller harder-tosee zooplankton become more common which favors jellyfish over fish predators because jellyfish hunt by touch not sight. Jellyfish populations Because they grow quickly jellyfish blooms have become widespread. Such blooms have led to the fish farms being wiped out by jellyfish, widespread beach closures, and the collapse of fisheries in the Black Sea. Jellyfish populations There is considerable concern that increasing jellyfish populations may result in permanently changed marine ecosystems. With populations of their predators reduced, larger jellyfish populations because they consume so many larval fish and eggs, may suppress the recovery of fish stocks permanently. Polyps Most polyps have tubular bodies and there is a mouth surrounded by tentacles. The tentacles capture prey which is then transferred to the gastrovascular cavity and digested there. Polyps are sessile and are attached to the substratum by a pedal disk. Youtube Anemones http://www.youtube.com/watch?v=O7_Is X-WZoc Life Cycles In cnidarian life cycles polyps and medusae play different roles. Life cycles differ among groups, but usually a zygote develops into a planula larva, which is free-swimming. This larva settles and develops into a polyp. Life Cycles The polyp may reproduce asexually and generate other polyps. Or as in the Hydrozoa (hydras) and Scyphozoa (jellyfish) polyps bud off medusae. These medusae are generated asexually, but each medusa is either male or female and produces gametes, which are shed into the water and produce zygotes beginning the life cycle again. Figure 13.09 7.9 Life cycle of Obelia, a marine hydroid. Life Cycles In the Anthozoa (sea anemones and corals) there is no medusa stage and all individuals are polyps. Both asexual and sexual reproduction take place in Anthozoa. Gametes are produced, but new individuals can be budded off too. Classes of Cnidaria There are 4 classes of Cnidarians Class Hydrozoa: Hydras, colonial hydrozoans inc. Portuguese man-of-war. Class Scyphozoa: most of the larger jellyfish Class Anthozoa: Sea anemones, hard corals, sea fans and sea pens Class Cubozoa: box jellyfish. Small group once considered an order of Scyphozoa Hydrozoa Most Hydrozoa are marine and colonial, but Hydra a freshwater hydrozoan is common in the U.S. and often seen in biology classes. Hydra is only about one inch long and has typical polyp form. It has no medusoid stage. Pink-hearted hydroid (Hydrozoa) Porpita (a colonial Hydrozoan) Hydra (Hydrozoa) Colonial Hydrozoa Individual polyps in colonial forms are referred to as zooids and specialize in particular tasks. The commonest are feeding polyps: gastrozooids. These capture and partially digest prey before emptying the food into the common gastrovascular cavity. Colonial Hydrozoa In colonial Hydrozoa the epidermis, mesoglea, and gastrodermis are all continuous making it difficult to tell where one individual ends and the next begins. Most colonial Hydrozoa are also surrounded by a non-living supportive protein-chitin envelope secreted by the epidermis and called the perisarc. Figure 13.09 7.9 Life cycle of Obelia, a marine hydroid. Colonial Hydrozoa Reproductive polyps (gonangia) produce medusae that leave the colony and produce gametes. These medusae are usually quite small (no bigger than a few cm). Unlike scypohozoan medusae the edge of the bell projects inwards forming a lip or shelf called a velum, which reduces the size of the opening of the bell. Muscular contractions and relaxations alternately fill and empty the bell moving the animal by a form of jet propulsion. Colonial Hydrozoa: Physalia Some hydrozoans known as siphonophores (including the well-known Physalia the Portuguese man-of-war) form floating colonies. The colony includes several forms of modified medusae and polyps. Colonial Hydrozoa: Physalia Physalia has a large float (a modified polyp) which is filled with carbon dioxide and this acts as a sail. There are multiple different polyps that hang beneath the float including feeding polyps, reproductive polyps and long stinging tentacles. Figure 13.14 Physalia Portugese Man-of-War Fig 7.12 Scyphozoa The class Scyphozoa includes most of the large jellyfish and the medusa is the dominant life stage. They are entirely marine. Most are 2-40 cm in diameter, but a few species (including Cyanea next slide) may be 2 meters in diameter with 60 meter tentacles. Scyphozoans bells differ from those of Hydrozoans in that they do not have a velum. Scyphozoa Bells vary in shape from helmet-like to shallow saucers. Many bells have a scalloped edge and the notches contain sense organs called rhopalia. Rhopalia include statocysts that assist with balance, other sensory cells that sense chemicals and in some cases simple eyes (called ocelli) Figure 13.16 Giant Jellyfish Cyanea capillata Fig 7.14 Scyphozoa Most Scyphozoans are pelagic, but in one unusual order the medusa attaches to seaweed. Figure 13.19 Thaumatoscyphus hexaradiatus sessile medusa Scyphozoa Scyphozoans have a typical medusa with a large bell and long tentacles. The mesoglea is thick (hence the name jellyfish). They feed on all sorts of small animals from protozoa to fish, which are stung and captured by the tentacles and transferred to the gastrovascular cavity. The gastrovascular cavity is complex in structure with 4 gastric pouches that connect with a series of radial canals and join a ring canal that run around the outside of the bell. The complex gastrovascular cavity allows nutrients to circulate around the whole animal. Figure 13.17 Moon Jellyfish Aurelia aurita Gastric pouches Radial canals Ring canal Scyphozoa In Scyphozoa the sexes are separate (gonads are located in the gastric pouches) and fertilization occurs inside the gastric pouch . Zygotes may be brooded or released into the water and they develop into a ciliated planula larva. The planula larva attaches to a substrate and develops in a series of stages into a strobila, which buds a medusa-like ephydra that grows into an adult medusa. Figure 13.18 Aurelia (moon jellies; Scyphozoa) Lion’s Mane jellyfish (Scyphozoa) Cubozoa The cubozoans are a small entirely marine group in which the medusa is the dominant form (polyps are either inconspicuous or unknown) The bell of cubozoans is almost square in section (hence “cube” ozoans and box jellyfish), it is not scalloped, and there is a flattened structure at the base of each tentacle called a pedalium, which facilitates identification Box jellyfishes (Cubozoa) Pedalium Cubozoa Cubozoans are very strong swimmers (like Hydrozoans they possess a velum-like structure called a velarium that enhances propulsion) and they are very effective predators, mainly on fish. They produce highly toxic venom and often have very long tentacles. Their rhopalia each contain 6 eyes as well as other sense organs. Chironex fleckeri (box jellyfish) has tentacles that can be 10 feet in length More than 100 people have died from stings in the past century in northern Australia. Box jellyfish video http://www.youtube.com/watch?v=uIf0kR pkQ_0 Anthozoa Anthozoa (“flower animals”) are polyps with a flowerlike appearance. They are entirely marine and there is no medusa stage. The Anthozoa includes three groups: Zoantharia: sea anemones, hard corals Octocorallia: sea fans, sea pansies, sea pens, soft corals Ceriantipatharia: tube anemones and thorny corals. A small group with few species. Orange sea pen (Octocorallia) Sea anemones (Zoantharia) Figure 13.21 Anthozoa: Zoantharia: sea anemones and hard corals Have a hexaramously symmetrical bodyplan (based on multiples of 6). In contrast octocorallians are based on multiples of 8. Tentacles are simple tubular structures in contrast to octocorallians which have featherlike tentacles. Anthozoa: Zoantharia: sea anemones and hard corals Sea anemone polyps are much larger and heavier than hydrozoan polyps. Usually they are colorful and may be up to 4 inches in diameter. Sea anemones are cylindrical with a crown of tentacles arranged in one or more rings around the mouth. Figure 13.21 Fig 7.19 Sea anemones Anthozoa: Zoantharia: sea anemones and hard corals Sea anemones are carnivorous and feed on fish or any other suitably sized prey. Sea anemones depending on the species may be hermaphroditic or have separate sexes. Zygote develops into a ciliated larva that settles and becomes a polyp. Asexual reproduction by fission also occurs. Anthozoa: Zoantharia: sea anemones and hard corals Hard corals (or scleractinian corals) are effectively miniature sea anemones that live in calcareous cups that they themselves secrete from their epidermis. The cup is made of calcium carbonate (CaCO3) and the coral can retreat into it when threatened. Figure 13.27 Hard coral polyp Anthozoa: Zoantharia: sea anemones and hard corals When contracted most corals are very difficult for fish or other predators to extract. The skeleton is secreted entirely below the living tissue and so is an exoskeleton. Coral polyps Anthozoa: Zoantharia: sea anemones and hard corals In colonial corals the skeleton may become massive over the years with living coral occupying only a thin sheath of tissue on the surface. The gastrovascular cavities of polyps are all connected through this tissue. Anthozoa: Zoantharia: sea anemones and hard corals The patterns formed in coral rock are caused by the growth patterns of the coral and the arrangement of polyps. For example, in brain corals, the polyps are arranged in rows. The rows are well separated, but the polyps that make up each row are very close together and their cups merge. As a result, the skeleton of the colony looks like a human brain with valleys separated by ridges. Brain coral (Anthozoa) Anthozoa: Octocorallia Octocorals all have 8 pinnate (feathery) tentacles and include soft corals, sea pens, sea fans, and gorgonians. They are colonies of polyps connected to each other by a coenchyme (consisting of mesoglea, spicules and connecting tubes) and forming elaborate branching structures. The gastrovascular cavities of polyps in colonies are connected by tubes called solenia. The solenia run through a gelatinous mesoglea, which is enclosed by epidermis. Figure 13.31 Polyps of an octocorallian coral 7.26 Anthozoa: Octocorallia Octocorals have an endoskeleton in which stiffening elements are secreted into the mesoglea. In some there is a central supporting rod of protein (gorgonin) or calacareous spicules that runs through the coenchyme. (Coral jewelry is made from such rods taken from the red coral Corallium.) Anthozoa: Octocorallia The coupling of spicules (sometimes fused together) with the stiff, but flexible protein gorgonin (similar to keratin) provides enough structural support for large fanlike or branched colonies of octocorals to develop. Figure 13.33b Sea fan (gorgonian) Figure 13.33a Red gorgonian Soft corals Some octocorallians lack the central supporting rods and are very soft. These soft corals have fleshy bodies that contain calcareous spicules in the mesoglea. Figure 13.32 7.27 Soft coral (Octocorallia) Anthozoa: Octocorallia Octocoral colonies may be in the form of mats or ribbons, feather-like or clusters of vertical branches. Often they are brightly colored: red, orange, yellow or purple. Figure 13.22a Orange sea pen (Octocorallia) Fig 7.20 Coral Reefs Coral reefs are found in shallow waters in the tropics. They are calcareous structures and what makes them unique as geological structures is that they are formed by some of the organisms that live on them, specifically reef-building corals and coralline algae. They are the largest living structures on the planet. Coral Reefs Reef-building corals contain symbiotic algae (zooxanthellae) that supply a significant part of the coral’s energy in exchange for protection and access to light. These algae require light for photosynthesis and so reef-building corals can live only in clear waters less than 100m deep (and most species occur in much shallower waters). Coral Reefs Many other cnidarians on reefs including octocorallians, sea anemones and hydrozoan corals also have zooxanthellae and are similarly restricted in their vertical distribution. Coral Reefs In addition to light, reef building corals require warm water where the average minimum temperature is at least 20ºC. As a result of their narrow tolerances, coral reefs are found only in waters between 30º N and 30º S of the equator. Coral Reefs Coral reefs are restricted to the Caribbean, Indian Ocean and tropical Pacific. Because of their narrow tolerances coral reefs are absent from much of the Atlantic. Water tubidity, because of the sediment carried by large rivers limits corals along the east coast of South America and west coast of Africa (cold water currents also restrict corals off Africa). Types of Coral Reefs Three general types of coral reef can be recognized. Fringing reefs: are the commonest type and project into the sea directly from the shore. Barrier reefs: are separated from adjacent land by a lagoon. Great Barrier Reef is the longest at >1000 miles on NE coast of Australia. Atolls: rest on summits of submerged volcanoes. Usually circular/oval with a central lagoon. Parts of the reef platform may emerge as islands. Shumann Island, Papua New Guinea (fringing reef). Pohnpei Atoll Micronesia Coral Reefs A reef platform (the layer of coral rock) may extend considerably below the current photic zone (reefs more than 1000m deep are known). How is this possible? Growth of reef platforms occurred as a result of changes in sea level or subsidence of the substratum. Coral Reefs In the late Pleistocene sea levels were about 120m below current levels when lots of water was locked up in ice sheets. As the Earth warmed between 18,000 and 7,000 years ago sea level increased by about 1cm/year (very fast). Corals simply grew upwards as the sea levels rose. Coral Reefs Reef platforms of great thickness are due to subsidence. Most atolls sit over volcanic seamounts that have subsided and as they have sunk corals have kept up with rate of subsidence. Charles Darwin was the first to figure this out. Coral Reef Zonation Coral reefs show considerable zonation depending on exposure to wave action. The seaward side of reef rises from the depths to just below surface and may be gently or steeply sloped. Large domed or columnar corals occur between 10 and 60m depth. Usually the reef front is not a smooth wall but rather a series of finger-like projections. This pattern disperses wave energy. Coral Reef Zonation The reef crest is where the reef approaches closest to the surface. There is high wave stress here and corals such as elkhorn corals predominate. Behind the reef crest is a reef flat which is quite protected and contains smaller, delicate branching corals. Figure 13.34 13.34 Coral Reef Diversity The waters in which corals are found are nutrient poor. The clear, blue color of the water is a tip-off to this. Productive waters have a lot of phytoplankton. As a result, they are usually green and light is absorbed quickly. Coral Reef Diversity Despite being in nutrient poor water, coral reef ecosystems are some of the most productive marine environments. This is because the populations of algae and symbiotic zooxanthellae carry out a huge amount of photosynthesis and so form the basis for an extensive food web. Coral Reef Diversity Besides cnidarians large numbers of sponges, molluscs, clams, tunicates, and bryozoans live on the reef. In addition, sponges, clams, and some worms bore into exposed coral. Coral Reef Diversity The huge numbers of holes and crevices offer shelter to shrimps, crabs, worms, molluscs, fish and other animals. All of these smaller animals attract large number of predators including fish, turtles, and sharks. nudibranch Puffer fish Blenny Green Sea turtle Coral reef, Indonesia Schooling jack fish Nature Conservancy Video clips http://www.nature.org/joinanddonate/resc uereef/explore/video.html Kimbe Bay Belize fish spawning Threats to coral reefs There are numerous threats to coral reefs. These include nutrient enrichment from sewage and agricultural runoff and overfishing of herbivorous fish, which result in heavy algal growth. In addition, sediment resulting from deforestation reduces water clarity and covers corals. Threats to coral reefs Global warming from increased levels of carbon dioxide also threatens reefs because when water becomes too warm corals expel their zooxanthellae (coral bleaching) and die. In addition, higher carbon dioxide levels in the atmosphere are lowering marine pH levels making the water more acidic. This makes it harder for corals to produce calcium carbonate and may also dissolve corals. Nature Conservancy Video clips http://www.nature.org/joinanddonate/resc uereef/explore/video.html Palau Youtube corals reefs Truk Lagoon 2:30 http://www.youtube.com/watch?v=ju6QxK6FJM&mode=related&search= Coral reef, Komodo Island National Park, Indonesia Phylum Ctenophora The phylum Ctenophora is a small phylum of fewer than 100 species of comb jellies all of which are marine. Comb jellies are named for the 8 rows of short, comb-like plates of long cilia they beat in order to move. Most ctenophores are free swimming. Beating of the cilia in each row begins at the aboral end and all plates beat in synchrony to move the ctenophore. An organ called the apical sense organ coordinates beating of the comb rows. Figure 13.35a 7.28 7.30 Figure 13.35b Phylum Ctenophora Ctenophores are quite similar to cnidarians in many ways: but there are a number of differences: Similarities: both have: nerve net diploblastic with thick gelatinous mesoglea Pelagic, transparent floating predators, slow moving Single oral cavity Tentacles solid not hollow Differences: Biradially symmetrical rather than radially symmetrical. Ctenophores lack nematocysts, have colloblasts Ctenophore cells are multiciliated Cnidaria swim by jet propulsion, ctenophores by beating of combs Like cnidarians ctenohores have no anus but possess anal pores, small openings to the outside from the gastrovascular cavity. Figure 13.36a Phylum Ctenophora Many ctenophores possess two long tentacles that are covered with adhesive cells called colloblasts, not nematocysts as in the cnidarians. However, one species of ctenophore does carry nematocysts, which it appears to obtain from cnidarians it eats. Unlike cnidarian tentacles, those of ctenophores can be retracted into pits or sheaths. In ctenophores without long tentacles the body is covered with colloblasts and the whole surface is used to trap prey. Small tentacles transfer prey to the mouth. http://www.marlin.ac.uk/images/taxonomy_descriptions/Ctenophora.jpg Phylum Ctenophora Ctenophores can be major predators of larval fish and other zooplankton such as crustaceans. The introduction of Mnemiosis leidyi an invasive species of comb jelly into the Black Sea about 25 years ago caused the collapse of the local anchovy fishery. The comb jellies consumed fish eggs and larvae as well as competed with fish for zoo plankton Figure 13.38 Ctenophore video clips http://www.oceanfootage.com/stockfoota ge/Comb_Jelly_Footage/owner%3Dekovac s//?DVfSESSCKIE=8cdc73b166081cdf4f03 3fe0c520befaa7ddb171