["Shape-shifting Spreading the arms helps Like other soft-bodied octopus the connecting webbing species, the day octopus can cast a larger shadow change its shape along with its color. By spreading its body and casting a shadow over lurking prey, it may encourage a shade- loving crab to venture farther from its hiding place. Slit-shaped pupil splits light into its constituent wavelengths, enabling the octopus to detect different colors, even though its eyes lack the color-sensing cells found in other animals with color vision False eye spots may direct predators away from the most vulnerable body parts, but their exact function is unknown 150 \u2022 151 coral reefs Interbrachial web\u2014a stretch of skin connecting the base of the eight arms\u2014is packed with color-changing cells (chromatophores) changing color Where sunshine penetrates the sea, many animals flaunt colors, and some even change color when their behavior alters. The switch can occur rapidly because it is controlled by lightning-fast nerve impulses. Cephalopods, such as octopuses and cuttlefish, are among the most impressive color changers. Within seconds, an octopus sensing danger can blend in with the seabed and vanish or, spotting a potential mate, impress with bold patterns.","","Marine science for the masses An 1850s issue of the popular illustrated Parisian magazine Le Magasin Pittoresque includes a delicate wood engraving of named species of bivalves, sea snails, and a sea slug in a project to educate the wider population about marine life. the ocean in art art meets science Natural history illustration came of age in the 19th century in response to 152 \u2022 153 coral reefs public fascination with voyages to formerly remote regions and a scientific need for accurate representation of specimens. The sea was a new frontier for exploration, and the complexity and vibrancy of its myriad life forms was both a challenge and an inspiration to the artists recording them. Describing sea creatures was historically Britain\u2019s HMS Challenger returned from challenging, as specimens rapidly lost their a 4-year voyage around the world in the colors and shrank after being preserved 1870s, with samples trawled from many in alcohol. The Zoological Society of different ocean depths and new data on London\u2019s solution was to use art painted the chemistry of seawater, currents, sea at the source. From 1812, tea merchant temperatures, and the geology of the John Reeves spent 19 years in Macao, seafloor. One of many enthusiasts fired where he commissioned Chinese artists up by the findings was German doctor and to create detailed paintings of fish. Many zoologist Ernst Haeckel. His major work of these paintings became iconotypes: the Kunstformen der Natur features 100 stunning basis for the description of a new species. lithographs that reveal the symmetry and beauty of the ocean\u2019s smallest life forms. Nudibranchs (1899\u20131904) Crossing the boundaries between scientific A ground-breaking 1855 study of the illustration and art, Ernst Haeckel\u2019s nudibranchs Atlantic by US naval officer Matthew (soft-bodied marine gastropods) are remarkable Fontaine Maury inspired naturalist James for their detail and harmonious arrangement. M. Sommerville to commission a dazzling watercolor illustration of its species. Ocean Life (1859) The grouping may be considered, rather as An idealized assembly of 75 vibrant an intellectual truth than as a literal view. It sea creatures and corals from a variety might be termed a mental oasis of the sea \u2026 of geographical zones was created by physician and naturalist James M. JAMES M. SOMMERVILLE, INTRODUCTION TO OCEAN LIFE, 1859 Sommerville to illustrate his pamphlet Ocean Life. Art professor Christian Schussele\u2019s watercolor for the lithograph has all the hallmarks of a still life painting.","cleaning hosts Cleaner shrimp may enter the Sunlit coral reefs are fertile, physically complex places with a rich host\u2019s mouth biodiversity. As a result, more species evolve to be specialized to pick food from in what they do, reducing competition in such a crowded habitat, its jaws and teeth and many are driven into close relationships. Shrimp that once scavenged for food now pick parasites and dead skin off living Cleaning the hunter hosts and even advertise their services at cleaning stations. In this Some cleaner shrimp form strong bonds mutually beneficial symbiosis, visiting fish \u201cclients\u201d are cleared with moray eels. By sharing the eel\u2019s lair, of potentially harmful pests, while the cleaner shrimp get a meal. they get some protection from predators. coral reefs 154 \u2022 155 Colorful cleaner Long white antennae may be On coral reefs in the Indo-Pacific, the distinctive waved around as part of the visual red-and-white pattern of the scarlet skunk shrimp signal that advertises the shrimp\u2019s (Lysmata amboinensis)\u2014named for its stripes\u2014may cleaning behavior to a \u201cclient\u201d fish serve as a visual signal to fish at cleaning stations. Several species of skunk shrimp act as efficient fish cleaners in tropical waters around the world. Red stripes are caused by a chemical pigment called astaxanthin in the crustacean\u2019s carapace","CLEANER FISH Fish from groups as diverse as gobies and wrasses have independently evolved cleaning behavior on coral reefs. Like cleaner shrimp, they consume skin parasites, and most share a similar striped color pattern, the stripes often being blue or dark. Their cleaning station is typically a conspicuous seabed feature, such as a coral outcrop. Here, \u201cclient\u201d fish gather and may solicit the attentions of waiting cleaners through their posture or swimming movements. BLUESTREAK CLEANER WRASSE (LABROIDES DIMIDIATUS) Large eyes are used to spot \u201cclients\u201d; in dim light, shrimp may be more wary of approaching potentially dangerous predators and exaggerate their precleaning signals, such as leg rocking and waving antennae Legs rock the body from side to side when approaching a shrimp-eating \u201cclient,\u201d helping reinforce the cleaning intent and avoid accidental attack by a predator","Under cover making a mask The spider decorator crab (Camposcia retusa) is so devoted to staying hidden Although crabs can famously use their pincers in self-defense, many that the only parts of its body not species adopt a less confrontational approach: they wear a disguise. covered by debris and living things are By plastering their body with flotsam or cuttings\u2014anything from those it uses to collect the masking scraps of seaweed to colonial animals like sponges\u2014they can blend material\u2014its pincers. in with the seabed. Much of what they wear keeps on growing, becoming a living mask that improves the deception over time. Slender pincers have sharp cutting edges for snipping pieces of algae or sponge Body debris includes fragments of seaweed, sponges, small stones, and shells collected by the crab","Hooked hairs grow in CRAB D\u00c9COR Anemones held Sponge in place bunches, which provides by pincers on crab\u2019s back a stronger grip when Some crab collectors are holding debris specialists. Those of the POM-POM CRAB SPONGE CRAB family Dromiidae camouflage Velcro crab themselves with sponges. The body of the spider They have hooked rear legs decorator crab is covered for transferring a sponge to the in hook-tipped hairs. The back of their shell. Pom-pom hooks work like Velcro, crabs (Lybia) even use d\u00e9cor securing objects to the with a sting: they carry tiny body transferred there anemones (Triactis) in their by the pincers. pincers to ward off predators. The legs, as well as the 156 \u2022 157 coral reefs body, are covered in hairs, so practically the entire crab can carry debris Yellow or red sponges stay alive and keep growing when attached to the crab Brightly colored sponges may produce noxious chemicals that repel any predators not fooled by the disguise","Fleshy projections (papillae) cover the body Anus serves as both an exit for excreted waste and an inlet for oxygenated water required for respiration coral reefs 158 \u2022 159 Sharing resources The thorny sea cucumber (Colochirus quadrangularis) lives in sunlit waters to Tentacles carrying food about 380 ft (115 m) deep. When feeding, are inserted one by one it rears upward and spreads its tentacles into the mouth to trap food suspended in the water. Some of the food may also be taken by Feeding tentacles tiny shrimp living in among the tentacles. The reef-dwelling sea apple (Pseudocolochirus seabed feeding violaceus) is a colorful sea cucumber that uses Animals that live sedentary or slow-moving lives on the seabed frilly tentacles to trap make the most of the food that comes to them. Suspension tiny particles, such feeders trap particles of living or dead material from the water as algae, zooplankton, using bristles or tentacles. Detritivores specialize in dead and dead material, from material, known as detritus, which can be trapped in the water sunlit shallow water. or gathered from the seabed. In deep water, detritus raining from above replaces sunlight as the main source of energy.","Body is flexible and muscular Feathery feeding tentacles trap particles of food WHALE FALL Scavengers eat Worms and Mussels, the soft tissue crustaceans clams, and tube Detritivores recycle matter and energy back into the food web. SCAVENGER STAGE Invertebrates worms drawn A dead whale first provides food for feed on the to bacteria mobile scavengers, including sharks remains and hagfish, then slower ones like Bacteria sea pigs (a type of sea cucumber), invade bones then colonists such as worms and crustaceans. When the flesh is gone, OPPORTUNISTIC STAGE SULFOPHILIC STAGE communities of bacteria in the bones produce hydrogen sulfide and use it in place of sunlight to fuel the manufacture of carbohydrate (see p.262).","Pigmented blue The mandarinfish (Synchiropus splendidus), found in shallow inshore reefs of the Western Pacific, is one of only two known vertebrate species whose blue color comes from a pigment rather than structural scattering of light. The striking blue coloration is used as a warning to predators that it is protected by a powerfully unpleasant-smelling toxic mucus secreted through the smooth, scaleless skin. coral reefs 160 \u2022 161 blue underwater Vivid colors of fish on a coral reef help signal species identity in this crowded community. Blue and yellow are the most common: they contrast well and transmit far underwater, while red is effective in shallower depths. Much of this color comes from pigments that absorb some wavelengths of light and reflect others. But some\u2014notably blue\u2014is structural. This happens when objects, such as microscopic crystals in a fish\u2019s skin, bend wavelengths to different degrees\u2014in the way raindrops produce a rainbow. PIGMENTARY AND Incoming light Blue light STRUCTURAL BLUES with various is reflected wavelengths Black, brown, yellow, and red pigments CYANOPHORE are produced in a fish\u2019s skin by cells called Shorter blue wavelengths chromatophores. So far, blue-pigmented Blue pigment are reflected chromatophores (or cyanophores) are absorbs all known only in the mandarinfish and a close IRIDOPHORE relative. Pigments absorb wave-lengths of wavelengths except blue light, except for the color that we see, which is reflected back. Other skin cells, called Crystal splits iridophores, contain guanine crystals. light into As light passes through the crystals, the different wavelengths bend to varying degrees. wavelengths Shorter blue wavelengths get scattered back and intensified, so the skin appears blue. This explains the blue color of Longer wavelengths other kinds of coral reef fish. are transmitted through the cell","","inflatable bodies GULPING WATER The related pufferfish, balloonfish, and porcupinefish all share the startling When frightened, pufferfish and porcupinefish defensive strategy of rapid inflation of the body to a near-spherical form in transform by rapidly swallowing water into the order to deter attacks from larger animals. They also produce potent toxins. stomach, which expands like a balloon, filling However, these defenses sometimes come too late\u2014large, quick predators all available space in the body cavity. Powerful may snap up a fish before it has fully inflated, which can result in the death sphincter muscles at either end of the stomach of both the fish and the predator. The unrelated swell sharks adopt a similar prevent liquid and gas passing into the gut during strategy, but they do not achieve the extreme roundness seen in the others. inflation. When the perceived threat has passed, the water is regurgitated or belched back out of the Difficult mouthful mouth and the fish deflates. The body of the long-spine porcupinefish (Diodon holocanthus) is covered in rows of scales that extend into Backbone Water flow needlelike spines. In a relaxed fish, the spines lie flat against the body, but when the fish swells, the skin tautens and the Stomach spines are pulled erect, increasing the deterrent to predators. UNINFLATED FISH Backbone INFLATED FISH Stomach inflates around the backbone coral reefs 162 \u2022 163 Uninflated fish uses its undulating pectoral fins to propel itself forward Skin contains collagen fibres, which help the skin to expand as the fish inflates Body inflates in 1\u20132 seconds (it takes slightly longer to deflate)","Blotchy body coloration serves as camouflage against seafloor Inflated fish struggles to swim Spines are modified bony scales","","Dugong Hunt (1948) In this bark painting of fishermen in northern Australia\u2019s Gulf of Carpentaria, a dugong\u2014a sea mammal especially valued by Aboriginal people\u2014is pulled from the depths. The work is attributed to the Aboriginal artist Jabarrgwa \u201cKneepad\u201d Wurrabadalumba. the ocean in art aboriginal sea stories The art of Aboriginal peoples in Arnhem Land, in Australia\u2019s Northern 164 \u2022 165 coral reefs Territory, reflects their ancient connection with animals in coastal waters and rivers. Traditionally, Aboriginal peoples use art (and oral literature) rather than the written word to pass on their knowledge and tell stories. Works may convey practical skills, such as fishing, or legends of cultural or spiritual significance, often using symbols passed down through millennia. Barramundi (20th century) Of all the fish featured in Aboriginal art, brush to create fine lines. In the northeast, Characteristic of western Arnhem Land art, a pair barramundi reign supreme. For millennia, the bark paintings of the Yol\u014bu people tell of barramundi are painted on stripped bark in the they were painted on rocks in sites that are tales of seafaring ancestors and legendary traditional X-ray style. The skeleton and organs now protected by the Kakadu National sea creatures and impart centuries of are mapped out inside the outline of the fish. Park. Today, the fish is widely reproduced knowledge as the caretakers of the coast. on paper and in bark paintings\u2014an art Dreamtime Spirits form that has become widespread in the After a barramundi fishing party and Fish (20th century) past 70 years. The bark is stripped from illegally encroached on a sacred site in The rock art sites in Kakadu National gum trees, then heated and flattened to 1996, Yol\u014bu artists in Yirrkala produced Park are decorated with paintings and create a surface for natural pigments. a set of paintings known as the Saltwater drawings dating back to c.26,000 bce, as Barks to explain their ancient traditions well as more recent works. As is typical In western Arnhem Land, subjects are and assert ancestral rights. In 2008, the of Aboriginal rock art, many artists have depicted in X-ray style (see left). In the east, High Court of Australia confirmed that retold stories by painting over older work. sea creatures and fishermen are rendered the area\u2019s Aboriginal owners have exclusive This painting of X-ray-style fish with with intricate cross-hatching, known as access rights to tidal waters along 80 per- ceremonial figures is one of 600 works raark, using reeds or human hair like a cent of the Northern Territory coastline. at 46 sites by the artist Najombolmi. By painting ... we are telling you a story ... We use our knowledge to paint from the ancient homelands to the bottom of the ocean. YOL\u014aU ELDER, SALTWATER: YIRRKALA PAINTINGS OF THE SEA COUNTRY, 1999","","Holding fire SECRETING MUCUS Cell Secreted Goblet cell The stinging cells (nematocysts) nucleus mucus produces and on anemone tentacles are usually The mucus secreted by various secretes mucus triggered by contact with another fish organs serves many purposes, not animal. To avoid being stung, the least acting as a barrier to pathogens. maroon clownfish (Premnas biaculeatus) Mucus contains a cocktail of secretes a body covering of mucus, carbohydrates and proteins, which to which the anemone does not react. varies according to the organ and species. Some evidence suggests that clownfish skin mucus appears to lack Basal lamina Proteins are produced, the proteins that anemones associate supports cells modified, and processed with potential threats or meals, and so for secretion in a series of sacs contact with the fish does not trigger a stinging response. MUCUS CELLS IN SKIN mutual protection 166 \u2022 167 coral reefs Several fish species live in mutually beneficial relationships with anemones, most notably clownfish. The fish gain protection from predators for themselves and their eggs among the anemones\u2019 stinging tentacles. The anemones benefit because the fish chase away anemone-eaters, drop food, and clear the anemones\u2019 tentacles of waste and debris. In addition, nutrients from the clownfish droppings boost the growth of algae (zooxanthellae), which live in even closer association inside the anemone, using light energy to make food. Larvae hatch a week after the eggs are fertilized by the male Safe nursery Clownfish embryos develop from eggs laid close to anemones, and the young fish are resistant to anemone stings from the moment of hatching. All clownfish begin life as males. The oldest and largest adults switch sex to become females.","sex changer Mostly yellow individuals are thought to be females Many fish change sex during their lifetime, which means that The wide gape an individual can breed as both a male and a female at is a threat different stages of its life. These fish are called sequential (or posture typical nonsimultaneous) hermaphrodites. Most of these sex changers, of moray eels including several species of wrasse, switch from females to males (protogyny). However, some species switch the other way The female stage (protandry), including the common clownfish and, according A major advantage of protandry, in which a male to some experts, the ribbon moray (Rhinomuraena quaesita). increases in size and becomes female, is that the larger females have greater bodily resources to invest in egg production, which requires a great deal of energy. coral reefs 168 \u2022 169 Watching and waiting Bright blue body All morays spend long periods hidden in coloration of male crevices with the head protruding. Relatively large eyes, expanded nostrils, and sensitive tentacles help the ribbon moray sense prey, which it seizes with its two sets of jaws\u2014one in the mouth, another in the throat. The ribbon moray is sometimes known as the leaf-nosed moray for its distinctive snout.","Nostrils flare into SIZE AND COLOR VARIATIONS delicate, trumpet- shaped structures Ribbon morays were once regarded as several species, because of their striking Tip of lower jaw color variation. However, some experts bears three short, think that they may develop from small black juveniles into blue adult males fleshy tentacles and then into large yellow females. Tall dorsal fin Typically less Can grow gives the body than 26 in up to 51 in (65 cm) long a ribbonlike (130 cm) appearance long Typically 26\u201337 in (65\u201395 cm) long JUVENILE ADULT ADULT MALE FEMALE The male stage As the ribbon moray matures, the jet black of the juvenile is thought to change to a distinctive, vivid electric-blue color. Current thinking suggests this may be the male stage of its life cycle. Bright blue coloration except on the dorsal fin, snout, and lower jaw, which are yellow","Single dorsal fin is deeply notched between the spines Large, bulging eyes provide almost 360\u00b0 field of vision Long, tweezerlike jaws; lower jaw projects slightly forward when fish feeds Bold patterns FEEDING ADAPTATIONS Picks plankton and pulses of from the water sound advertise The long, narrow jaws of the longnose territorial butterflyfish resemble a pair of Reaches deep boundaries bottlenose pliers, making them ideal into crevices for feeding in the crevices of the reef, Feeding in pairs where many other fish cannot reach. LONGNOSE Chaetodon lineolatus, which feed mainly on Some other butterflyfish, such as BUTTERFLYFISH coral polyps, tend to be monogamous; pairs bannerfish, have shorter jaws and work together to defend a feeding territory. bristlelike teeth, which are useful for snatching small prey, such as free-floating plankton, from the water, or picking food off the surface of the reef. BANNERFISH","Striking colors fade at night to make resting fish less conspicuous reef feeders Dancing color Butterflyfish are named for their colorful The 5,000 or more species of fish that live on coral reefs feed in many appearance and the way they flutter above different ways. This helps them divide up the available resources and the reef in \u201cclouds.\u201d The yellow longnose avoid competition. Only about 130 eat coral polyps directly, while butterflyfish (Forcipiger flavissimus, opposite) many more take tiny invertebrates hidden among the coral rock. and the smaller copperband butterflyfish Dominant among the polyp-eaters and rock-probers are the (Chelmon rostratus, above) are both found butterflyfish. Their tiny protractile mouths, armed with comblike teeth, in reefs in the Indian and Pacific Oceans. are perfect for foraging among crevices or rubble, from where they extract their miniature prey.","Spines of elegant Dorsal fin is yellow in Naso elegans unicornfish are fixed and mostly black in the otherwise in an open position, almost identical N. lituratus while those of most other surgeonfish are retractable Sharp end Like its close relatives, the elegant unicornfish (Naso elegans) possesses at least two forward- pointing spines on either side of the tail. A flick of the tail can inflict serious wounds. coral reefs 172 \u2022 173 Sharp spines at base of tail project outward from body Body coloration brightens with age: young specimens are gray, while older individuals have a yellow chest bladed fish Most fish are covered in scales, which are small, rigid plates that grow out of the skin. While all scales offer some form of protection, they vary greatly in structure and composition. Some groups have evolved modified scales that serve a particular function. Surgeonfish (also referred to as tangs or unicornfish) are distinguished by having large scales on the tail, modified into sharp spines, or scalpel-like blades. These usually lie flat, and the fish raises them when disturbed; however, in some species, they are fixed in a raised position.","Defensive weapons Like all surgeonfish, the elegant unicornfish (Naso elegans), sometimes called the lipstick tang, uses its spines as a form of defense. It is rarely aggressive toward other fish and is quite confident, perhaps because of its built-in weapons. Large eyes provide excellent color vision Small mouth, with orange \u201clips,\u201d contains a single row of sharp teeth used to shred seaweed","Small prey species Translucent cryptobenthic Long, tapered These fish, especially the hidden bottom-dwellers fish that is no more than body can be 2 in (cryptobenthics), are the some of the smallest and most 1 in (2.5 cm) long (5 cm) in length plentiful and account for almost 60 percent of fish consumed by larger species. Cryptobenthic larvae remain MAGENTA DOTTYBACK close to their parents\u2019 reefs, increasing their chances of Pictichromis porphyrea survival, so the fish supplies are constantly replenished. Laterally compressed URCHIN CLINGFISH Skin is smooth GLASS GOBY shape enables fish to Diademichthys lineatus and scale-less but Coryphopterus hyalinus swim through narrow is protected by a gaps in the reefs layer of mucus Male has bright red dorsal fin that it erects Foragers during courtship The water column zone of a reef sees slightly larger fish that feed on the smaller bottom-dwellers, coral polyps, plankton, and micro-invertebrates. They also clean the reef and some\u2014for example, the surgeonfish\u2014also consume algae, which helps ensure sunlight can reach the coral for photosynthesis. BULLSEYE CARDINALFISH Head has distinctive FLAME ANTHIUS COPPERBANDED BUTTERFLYFISH Ostorhinchus fleurieu dark stripe on the Pseudanthias ignitus Chelmon rostratus snout and electric blue lines at the eyes Snout is long and pointed with large mouth; upper Visiting predators Venomous dorsal The largest and most active predators of the reef\u2019s food spines are raised canine teeth are sometimes chain are the visitors that feed voraciously on fish, to deter would-be visible when mouth is closed crustaceans, and plankton, as well as benthic organisms predators such as algae, echinoderms, mollusks, tunicates, sponges, and hydrozoans. They range in size from the larger sharks and rays to huge schools of smaller species such as triggerfish and snappers. ORANGE-LINED TRIGGERFISH Enlarged scales above RED LIONFISH SCHOOLMASTER SNAPPER Balistapus undulatus the base of pectoral fin, Pterois volitans Lutjanus apodus behind the gill, are used to produce sound","Lower jaw carries a pair reef fish 174 \u2022 175 coral reefs of venomous canines used in defense Coral reefs cover scarcely 0.1 percent of the world\u2019s oceans but are home to around one third of all described species of marine STRIPED BLENNY fish. The three-dimensional structure of a reef\u2014as complex as Meiacanthus grammistes a rainforest on land\u2014has provided a rich variety of opportunities for fish to evolve in a multitude of different ways, helping them Long, sharp coexist and interact with the other life around them. preopercular spine protrudes between Color-changing reef dweller cheek and gill cover Streaked with blue over a pale brown ground, the elongate surgeonfish (Acanthurus mata) SABRE SQUIRRELFISH is capable of changing the background color Sargocentron spiniferum to dark brown. The species inhabits steep slopes and rocky bottoms and is often seen in large schools. This surgeonfish feeds almost entirely on zooplankton and can live for more than 20 years. Body can be up to 20 in (50 cm) long Sharp, lancelike defensive spines are folded into small horizontal grooves on each side of the tail Upper jaw has outer Jaws of adult fish row of strong canines have 26 small close-set and an inner band of teeth at the top and small teeth 24 at the bottom BLUEFIN TREVALLY Pectoral fins control Caranx melampygus swimming movement, stabilizing the laterally compressed body","Fleshy tassels (cirri) may By staying motionless mimic the white polyps that the pipefish perfects the project from a gorgonian coral deception cryptic fish Perfect match The red, elongated body and Coral reefs and shallow seas are visually complex spreading fins of a harlequin environments, and many species take advantage ghost pipefish bear a very of the intricate mix of color and texture to hide from close resemblance to the predators and prey. Ghost pipefish are true masters branching colony of a of the art of camouflage and crypsis, with different gorgonian coral. species specializing in mimicking the appearance and behavior of diverse reef-dwelling organisms.","Head down A head-standing posture may help a harlequin ghost pipefish blend in with vertical branches of coral. Other species of ghost pipefish adopt a similar behavior when hiding in seagrass. Relatively long, Eyes on side of head 176 \u2022 177 coral reefs pipettelike snout rotate in sockets, providing all-around vision even while the fish is motionless Tiny, toothless mouth at the tip of the snout sucks in prey (typically tiny crustaceans) Color shift The harlequin ghost pipefish (Solenostomus paradoxus) changes its appearance to be as inconspicuous as possible at each stage of its life. The transparent larvae start off in open water, where they live unobtrusively among the plankton. On maturity they live on the reef, where they are surrounded by gorgonian coral and crinoids and adopt strong colors and textures and a ragged outline.","specialized fins First dorsal fin, containing three About 99 percent of all bony fish, of which there are around 30,000 species, spines, is raised are classified as ray-finned fish; this makes them the most species-rich class of vertebrates on Earth. All members of this group have fins made up of in self-defense fine webs of skin supported by bony rays. The adaptability of this basic structure contributes to the group\u2019s success, with size, shape, and function varying to give fish the specialist abilities that set species apart. In triggerfish, the first dorsal fin is small and triangular and contains three thick, spinelike rays. If the fish is relaxed, the spines lie flush with the back, but if it feels threatened, it raises its spines so that they are perpendicular to the body. LOCKING MECHANISM Dual-purpose spine When the reef-dwelling clown If a triggerfish feels threatened, it raises triggerfish (Balistoides conspicillum) the spines in its first dorsal fin. The smaller is targeted as prey by larger species, secondary spine then slots into a groove on the lockable spine is an important the back of the larger front spine, locking it defense mechanism. In addition in place. The front spine cannot be lowered to making the triggerfish difficult until the secondary spine has folded back. to swallow, it can be used to wedge the fish in a crevice where the Front spine is locked predator cannot reach it. in upright position Second spine acts as trigger Both spines fold down together DORSAL FIN SPINES OF A TRIGGERFISH","Second dorsal fin ripples to provide source of propulsion Caudal fin used in short bursts of fast swimming\u2014 for example, when the fish flees a predator 178 \u2022 179 coral reefs Anal fins help propel fish through water Pelvic fins reduced to a pair of stout ventral spikes","","Venus in a half-shell (1st century ce) In the central peristyle (courtyard) of the House of Venus Marina, in Pompeii, a dazzling fresco panel of Venus takes center stage. Rising fully formed from her half-shell, she is carried forward by the wind caught in her red mantle and is attended by sea nymphs and a dolphin. the ocean in art seas of antiquity The sea played a key role in the two ancient civilizations that underpinned 180 \u2022 181 coral reefs much of Western culture. The city-states of ancient Greece fringed the Aegean and extended along the Mediterranean coast until they came under Roman rule in 146 bce. At its peak, the Roman Empire controlled all the lands around the Mediterranean, and many beyond. For both civilizations, the sea was a major source of inspiration in their mythology and art. An ocean floor (1st century ce) The perfection of classical Greek art became a melting pot for the cultures that A floor mosaic from a Roman calderium (hot bath) survives in statues, sculpted friezes, and preceded it. Looted classical treasures, features African fishermen chasing fish and sea mosaics but rarely in paint. The few such as bronze statues and paintings monsters. It was recovered from the House of frescoes that remain comprise painted of mythological scenes, were prized Menander in Pompeii. tombs in Paestum, southern Italy, dating and preserved through faithful copies. from the 5th century BCE, and fragments Hellenistic gods were renamed and their The sea personified (2nd\u20133rd century ce) from the much earlier Minoan civilization stories replicated in the frescoes and In Greco-Roman mythology, Pontus or Oceanus on the Greek island of Crete. Two in mosaics that adorned the homes and personified the sea and fathered a panoply of sea particular honor the sea: in Paestum, gardens of wealthy Romans. gods and sea life. In this detail from a Roman mosaic a lone diver flexes his body as he swoops in Utica, Tunisia, the watery god, with a beard of waves, down to meet the water; in Crete, Sea-life mosaics were favored for bath seaweed hair, and lobster-claw horns, is attended by dolphins, painted c.1500 BCE and restored houses and courtyard pools, especially erotes (winged cupids) riding on the backs of dolphins. in the early 20th century, play across the in the luxury villas of Pompeii, southern walls at the palace of Knossos. Italy. A famous mosaic from the House of the Faun celebrates the coastal bounty Further insights into the art of ancient of fish, lobsters, eels, and octopuses in a Greece have emerged through Roman art, design that was widely used. In coastal largely because the expanding empire colonies in Turkey and North Africa, mosaics featured sea gods and goddesses, (The dolphin) does not dread man, as though a such as Poseidon and Amphitrite, and stranger to him, but comes to meet ships, leaps and a host of fish-tailed mythical beasts. bounds to and fro \u2026 Dolphins were popular in mosaics PLINY, HISTORIA NATURALIS, 77\u201379 CE and frescoes as attendants to Aphrodite or playfully circling fishermen and ships. In his Historia Naturalis (77\u201379 CE), Pliny the Elder contributed to their magic with his tale of a dolphin who befriends a boy and carries him to school every day.","Jaws are covered with a mosaic of hard-wearing teeth, which are continually replaced throughout life TEETH AND DIET Blunt, fused teeth The jaws and teeth of fish show a huge variety of specialist forms HERBIVORE (QUEEN PARROTFISH) and functions. For example, the teeth of the queen parrotfish Sharp, (Scarus vetula), which is a herbivore, pointed teeth are completely blunt, with a highly CARNIVORE resilient, replaceable surface that (NORTHERN RED SNAPPER) is used to scrape the surface of rock and coral without damaging the underlying bone. In contrast, the carnivorous northern red snapper (Lutjanus campechanus), which feeds on small fish and crustaceans, has needle-sharp teeth that are used as weapons for stabbing and snagging its prey.","Excreting sand As the steephead parrotfish (Chlorurus microrhinos) feeds, it gouges particles of reef and rock from the substrate, which it ingests at the same time as its food. The particles pass through the gut undigested and are then excreted as fine white sand. Robust beak and powerful jaws are typical of excavating parrotfish reef scrapers 182 \u2022 183 coral reefs Lurid colors are indicative The sunlit surfaces of rocky and coral reefs make an ideal growing of males, but like most substrate for algae, which in turn provide an abundant food resource parrotfish, this species for other reef-dwellers such as parrotfish. These colorful fish, which changes sex from female are members of the wrasse family, have teeth and jaws that have evolved to male during its lifetime into a beaklike structure. Some species use their teeth to scrape algae from the reef surface, and in others, the beak is so tough that it can gouge into the reef. These so-called excavating parrotfish supplement their algal diet with animal matter in the form of coral polyps. Reef resources The ember parrotfish (Scarus rubroviolaceus, far left) and yellow-finned parrotfish (S. flavipectoralis, left) are closely associated with tropical reefs. They feed by scraping algae from the surface of rocks and corals using their beak. As well as providing a vast surface area for feeding, the complex reef structure also provides shelter for the fish.","sceoaasstal Rich in nutrients from ocean upwellings and land run-off, coastal seas have abundant plant and animal life. The shallow, light- drenched habitat is ideal for photosynthesis, especially for small plants and macroalgae rooted to the seabed.","","absorbing light Like all algae, seaweeds make food by photosynthesis, using sunlight\u2019s energy to convert carbon dioxide and water into sugar and other nutrients. All seaweeds use pigments to absorb this energy, while their fronds maximize the surface area for light capture. Different pigments absorb different color wavelengths present in white sunlight. Green chlorophyll predominates, but some seaweeds have accessory pigments that make them brown or red. The green color of the frond is due to the photosynthetic pigment chlorophyll, which is packed inside microscopic structures called chloroplasts coastal seas 186 \u2022 187 Selecting the spectrum The frond\u2019s cylindrical midrib Green seaweeds, such as sea grapes (Caulerpa has chloroplasts that carry out lentillifera), lack the accessory pigments of other photosynthesis; it transports colored seaweeds. Their color is due largely to the nutrients produced by chlorophyll, the same pigment used by land plants. photosynthesis to other parts Chlorophyll absorbs red and blue wavelengths of the seaweed of light\u2014for use in photosynthesis\u2014but reflects green wavelengths.","Grapelike ramulus stores sugars and other nutrients that are generated by photosynthesis; each frond consists of a branchlike midrib carrying rows of ramuli LIGHT AND HABITAT Red light Blue light penetrates penetrates Photosynthetic pigments more than help seaweeds thrive in less than 330 ft (100 m) different habitats. Blue and 165 ft (50 m) green light penetrate water Fronds are more deeply than other Deeper seaweeds have more pigment stiffened with wavelengths, enabling red for absorbing available wavelengths white mineral seaweeds that absorb these colors to grow better at SEAWEED DEPTH DISTRIBUTION Red seaweed greater depths. But the In addition to chlorophyll, quantity of pigment, as well red seaweeds\u2014such as this as the type, is important, Corallina\u2014have accessory too: some seaweeds of pigments known as other colors have higher phycobiliproteins, which pigment levels to absorb reflect more red light and more light in deeper water. absorb more green.","","spotlight species giant kelp Forming underwater \u201cforests\u201d rising more than 100 ft (30 m) from the seabed, giant kelp (Macrocystis pyrifera) is a type of algae that acquires food by photosynthesis. It is found off the North American Pacific coast from Canada to California and in temperate regions of all Southern Hemisphere oceans. Giant kelp is the largest living seaweed\u2014 Giant kelp anchors itself to the seafloor 188 \u2022 189 coastal seas the term for large algae growing on ocean using holdfasts\u2014rootlike attachment coastlines. They are supported by water, organs that form a tangled mass around which enable some seaweeds to grow to rocks, rubble, and debris on the seabed enormous lengths. Giant kelp is thought (see p.27)\u2014to prevent it from drifting to have the fastest linear growth rate on ashore. Rocky coastlines make ideal Earth and can increase its length by up fastening places. A primary stipe, or main to 2 ft (61 cm) per day. In clear, nutrient- stem, rises toward the surface, sending out rich waters where the sea temperature branching side stalks with leaflike blades, is 42\u201368\u02daF (5\u201320\u02daC), it can survive at each with a pneumatocyst, or gas bladder, depths of 175 ft (53 m), but it reproduces at its base that keeps it afloat. At the only if the water temperature is below surface, the kelp covers the water in a 65\u201368\u02daF (18\u201320\u02daC). type of canopy. Unless compromised by warming seas or overgrazed by sea A forest for carbon capture urchins, kelp can live up to 7 years. Giant kelp forests, like this one off the coast of California where blacksmith Kelp forests provide a key habitat damselfish shelter, absorb vast amounts for many animals. Holdfasts shelter of carbon dioxide. They act as natural invertebrates, fish and their larvae, and carbon sinks in a similar way to terrestrial shelter birds and sea otters during storms. forests and lower seawater acidity levels. Sharks, sea lions, and seals are also attracted to these rich hunting grounds. Pneumatocysts hold the blades upright, enabling kelp to take in sunlight for photosynthesis Staying afloat Balloonlike pneumatocysts are filled with a mixture of carbon dioxide, oxygen, and nitrogen that giant kelp cells both produce and absorb from surrounding seawater.","underwater meadows Asexual reproduction Turtle seagrass produces horizontal rhizomes that Most true plants cannot grow in the ocean. While the sunlit shallows can sprout plant clones, helping the species form be thick with seaweeds and other algae, these are solar-powered organisms extensive swards\u2014much like grasses do on land. that lack true leaves, roots, and flowers. However, seagrasses, such as turtle seagrass (Thalassia testudinum), are flowering plants. Rather than merely tolerating the salty ocean environment, they thrive in it; they bloom and set seed below the surface and cover vast areas of the seabed. At a time of global warming, these photosynthesizing green carpets are globally important as efficient stores of atmospheric carbon. coastal seas 190 \u2022 191","SEXUAL REPRODUCTION Sticky pollen bundles In turtle seagrass, genetic diversity comes from cross-fertilization Up to five Single during sexual reproduction. The flowers flower sexes are separate: each plant per plant produces either male or female per plant flowers. Male flowers release pollen at night in packages of MALE FEMALE Tropical seagrass carbohydrate-rich slime. The Turtle seagrass\u2014the favorite food of the slime attracts hungry invertebrates, Roots SEEDLING Seeds FRUIT green turtle (Chelonia mydas)\u2014is a common which help disperse the pollen to anchor sprout seagrass of the Caribbean. Neighboring female flowers. Fertilized flowers seedling to while fruit plants are interconnected clones, but they form seeds encased in small fruits. seabed is floating also reproduce sexually: their seed-packed The fruits float before settling on fruits disperse on the waves and establish the seabed, where each seed new colonies elsewhere. completes its germination.","producing light Colorless jelly Like many vulnerable jellyfish, the thumb-sized Many soft-bodied, jellylike animals of the open ocean can crystal jelly (Aequorea victoria), a hydrozoan produce light, a phenomenon known as bioluminescence. from the surface waters of the eastern Pacific Species as diverse as jellyfish, hydrozoans, and comb jellies turn Ocean, has a transparent body. This may help luminous if the water around them is agitated\u2014possibly to scare conceal it from predators against the open predators or attract tiny planktonic prey. The light, produced by water. Its light-producing photophores, which a chemical reaction in organs called photophores, is most striking are carried around the rim of its swimming when emitted in dark, deep water or at night nearer to the surface. bell, produce a momentary green glow. Rim of the bell contains photophores, but any light produced is only visible in dark conditions","Transparent bell pulsates to propel the crystal jelly through the water More than 100 tentacles hanging from the rim of the bell carry nematocysts (stinging cells); they immobilize and catch tiny planktonic prey HOW PHOTOPHORES WORK Light- Fluorescent Densely packed producer protein photophores glow Bioluminescence typically depends Photophores green and illuminate upon a light-producing chemical and around rim Enzyme the rim of the bell an enzyme. When stimulated by a trigger\u2014such as calcium, in the case 1. Enzyme locks on 3. Energized of the crystal jelly\u2014the enzyme to light-producer, protein emits makes the light-producer undergo causing it to emit green light a chemical change, which releases light. The distinctive green glow of energy as blue light 2. Fluorescent Glowing green the crystal jelly happens because protein absorbs Once activated, a cocktail of chemicals an additional component\u2014a light energy in the photophores of a crystal jelly fluorescent protein\u2014subsequently emits green light; each glow lasts for changes the wavelength of the just a few seconds. emitted light from blue to green. ARRANGEMENT AND FUNCTION OF CRYSTAL JELLY PHOTOPHORES","","alternating HYDROZOAN SEXUAL LIFE CYCLE generations The hydrozoan Obelia grows into an upright Animals that are attached to the substrate must get their branching colony of polyps, some of which nourishment close to the seabed. Hydrozoans\u2014relatives of generate pods that produce medusae. The anemones, corals, and jellyfish\u2014have polyps with tentacles free-swimming medusae release sperm and eggs that may rise only a short distance from where they are into the water. Larvae develop from the fertilized growing. But many also produce swimming, jellyfishlike bells, eggs and eventually settle on the seabed, where called medusae, as part of their life cycle. Medusae travel far they form new polyp colonies. The life cycles of in the water column, where they catch plankton beyond the other hydrozoans and related animals vary. Some reach of polyps. By alternating between polyp and medusa hydrozoans\u2014as well as corals and anemones\u2014 generations, food sources are more fully exploited. do not have a medusa stage, while certain jellyfish species exist only as medusae. Two generations Ernst Haeckel\u2019s Kunstformen der Natur (1904) depicts more Medusa swims than a dozen species of hydrozoan\u2014some as anemonelike in open ocean polyps (bottom) and others as medusae (top). Not all have a life cycle that switches between the two stages: some do not Pod become medusae and, like many other ocean invertebrates, releases experience open water only as microscopic larvae. medusae Medusa releases eggs and sperm Branching Fertilized colony egg Larva 194 \u2022 195 coastal seas OBELIA LIFE STAGES Polyps in close-up Tentacles trap The polyps of oaten pipes (Tubularia microscopic plankton indivisa)\u2014a European hydrozoan that and transfer them to the lacks a medusa stage\u2014carry rings of mouth at the center drooping tentacles on stems that extend up to 6 in (15 cm) from their base. Gonozooids are reproductive capsules that release larvae; the larvae develop directly into new polyp colonies","The Miracles of Mazu (18th century) An album of seven ink drawings, now housed in the Rijksmuseum, Amsterdam, Holland, describes the sea journey of a Song dynasty deputation to the Goryeo court in Korea in 1123 CE. The Chinese ships are under the protection of the sea goddess, Mazu. She is shown here in her traditional red robes, hovering on a cloud and carrying an ivory Hu tablet as a symbol of power.","The Story of the Great Flood (c.1870) In this depiction of a Hindu legend, Matsya (one of the 10 avatars of the Hindu God Vishnu) appears as half- human, half-fish. Here, together with seven great sages, he saves Manu (the first man) from the flood. the ocean in art asia\u2019s sea deities Thousands of years of art and storytelling reflect humans\u2019 longstanding awe 196 \u2022 197 coastal seas of the ocean. In most ancient cultures, the sea\u2019s unfathomable depths and unpredictability gave rise to numerous myths and beliefs, in which natural phenomena\u2014such as storms, currents, and fair winds\u2014were attributed to the whims and protective powers of sea gods and goddesses. In ancient Hindu texts (Vedas), Varuna goddess Mazu. Her story varies with the was the god of the sky, but in later tales, telling, but she was born Lin Moniang in he came to symbolize oceans, clouds, and 960 ce in a Chinese coastal town, where water. In Mysore painting, a classical art she was renowned for her swimming form of southern India featuring deities skills. She gained mystical status after she and mythological scenes, Varuna is often used trancelike powers to save male pictured riding the makara\u2014a hybrid sea members of her family from drowning at creature resembling a crocodile. Varuna sea. In 1281, she was given the official title also appears in miniatures that illustrate of Tianfei (\u201cPrincess of Heaven\u201d). the 5th-century bce Indian epic Ramayana, In the 1400s, Chinese admiral Zheng rising from the ocean to pacify Rama, an He had his great deeds carved in tablets of avatar of the god Vishnu. According to granite in two cities in China. He the legend, Rama\u2019s wife, Sita, was held attributed his seafaring success to Mazu\/ in captivity across the ocean, in Lanka. Tianfei, describing her as a \u201cdivine light\u201d In China, from the 11th century on, above the mast, calming the crew in rough the success of perilous sea journeys was seas. In reality, this light was probably St. often attributed to the beneficence of the Elmo\u2019s fire, a ball of light formed from electricity that appears on ships at sea in storms. The same goddess is evoked in an When we met danger, once we invoked the divine 18th-century album of ink drawings (see left), where she is pictured floating above name, her answer to our prayer was like an echo. Chinese ships, providing protection. Mazu worship spread to sailors and fishermen beyond China, and lacquered wooden statues, woodcut illustrations, and ADMIRAL ZHENG HE\u2019S DEDICATION TO MAZU\/TIANFEI, LIUJIAGANG TEMPLE, 1431 painted panels of the goddess are found in temples dedicated to her across the world.","Inner whorl of shorter labial tentacles transfers prey to the central mouth Tube is made from specialized stinging cells and sediment, all bound together by mucus","Outer whorl of longer Tube sanctuary marginal tentacles has Tube anemones cannot retract their stingers for catching prey tentacles, but when disturbed, they pull and self-defense themselves into their tube by contracting muscles along the length of their body. 198 \u2022 199 coastal seas The tentacles disappear as the anemone withdraws its body deep into the tube living in a tube Most anemones must make a living attached to a firm surface and can reach no higher than the stretch of their tentacles, but one group has evolved the means not only to live on soft sediment, but even to move up and down on the seabed. Tube anemones live inside vertical, partially buried, feltlike tubes. Instead of being anchored to the bottom, they can slip upward through the tube or sink deeper inside when danger threatens. Emerging tentacles A fireworks tube anemone (Pachycerianthus multiplicatus), from European coastal waters, erupts from the confines of its tube to sweep the water with its tentacles for planktonic prey. A mature specimen can have tentacles that span 12 in (30 cm). ANEMONE ANATOMY Mouth All anemones have just one opening Tentacle to their gut for ingesting food and eliminating waste. The gut is a simple Body Mesoglea cavity lined with digestive cells. Prey stalk (jelly layer) caught and paralyzed by the tentacles are transferred to the gut opening Digestive Epidermis before being digested by enzymes cells released from the gut lining. The basal disk is attached to a surface in Basal Gut cavity most anemones; in tube anemones, disk it is free to move up and down. CROSS-SECTION OF ANEMONE","Touch-sensitive antenna, Caruncle carries chemical one of three, aids navigation sensors and grows as an extension from the first when the fireworm crawls body segment over a reef Fleshy appendage, or parapodium, Fleshy projections carries sensory filaments called cirri called palps, one on and bundles of bristles known as chaetae; each body segment has either side of the a pair of parapodia antennae, are used to manipulate food Third, central antenna is shown here curving downward"]
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