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Fishes Chapter 24 I. Diversity  26,000 living species—more species than all other vertebrate groups combined  Adapted to live in a medium 800 x more dense than air  Can adjust to salt and water balance of environment  Gills extract oxygen from water that has 1/20 th the oxygen of air  Aquatic environment both shaped and constrained their evolution II. Ancestry and Evolution  A. Ancestor  Descended from freeswimming protochordate ancestor  B. Agnathans  Earliest fish-like vertebrates  Include extinct ostracoderms, and living lampreys and hagfishes  C. Placoderms  Fish with paired appendages and jaws that went extinct in Carboniferous with no living descendants D. & E.  D. Cartilaginous Fishes  Lost heavy armor and adopted cartilage as skeleton  Flourished during some periods, becoming nearly extinct during others  E. Acanthodians  Resemble bony fish but have heavy spines on all but caudal fin; sister group to bony fishes  Went extinct in lower Permian F. Bony Fishes  Dominant fishes today  2 distinct lineages—ray finned and lobe finned  Ray-finned radiated to form modern bony fishes  Lobe-finned include lungfishes, the coelacanth, and are sister group to tetrapods (amphibian ancestors) III. Superclass Agnatha: Jawless Fishes  A. Characteristics Lack jaws, internal ossification, scales, or paired limbs Pore-like gill openings and eel-like body B. Class Myxini: Hagfishes  43 species  Entirely marine  Scavengers and predators of annelids, molluscs, dead or dying fishes, etc  Nearly blind but locates food by acute sense of smell  Rasps hole into prey then eats from inside out with plate-like tongue  Glands secrete substance that becomes slimy in contact with seawater C. Class Cephalaspidomorphi  1. Diversity  41 species; 22 in North America and of these, half are non-parasitic brook variety  Marine lampreys can grow to length of 1 m.  All lampreys reproduce in freshwater streams, dying soon after 2. Parasitic Lampreys  Attach to fish by suckerlike mouth and sharp teeth rasp away flesh  Anticoagulant injected into wound to stimulate flow of blood  Wound may be fatal to host fish  Non-parasitic lampreys do not feed; digestive system degenerates and fish die after reproducing, within 2-3 years 3. Sea Lamprey Invasion  No lampreys existed in Great Lakes prior to 1829 when shipping canals were built  By the 1940’s, they existed in all the lakes  They decimated almost all fish species until populations finally declined due to lack of food and control measures IV. Class Chondrichthyes  A. Overview  850 species nearly all marine; 28 species live in freshwater  Ancient lineage but have survived due to well-developed sense organs and powerful jaws making them successful predators  Largest living vertebrates, after whales, reaching up to 12 m in length Whale shark reaches 43’ in length B. Subclass Elasmobranchii  1. Sharks A. Orders Carcharhiniformes—tiger and bull sharks which are coastal sharks and the hammerhead Lamniformes-- white and mako sharks which are large pelagic sharks Squaliformes—some of these are deep sea dwellers like dogfish sharks Orectolobiformes—carpet sharks like bamboo, nurse, and whale sharks Sharks More Sharks b. Outer Physiology  Streamlined fusiform body shape  Pointed nose with paired nostrils in front of ventral mouth; on hammerhead, nostrils on ends of “hammer”  Lateral eyes without lids  Tail has longer upper lobe (heterocercal)  Paired pectoral and pelvic fins, 1-2 dorsal fins, 1 caudal fin, and sometimes an anal fin  Tough, leathery skin with placoid scales that reduce water turbulence Body Structures of a Shark c. Senses  Olfactory organs can detect chemicals diluted 1/10 billionth their original concentration  Lateral line senses low frequency vibrations of prey over large distances  Excellent vision, even in dim water, used at close range  At close range, sharks are guided to prey by electric fields surrounding all animals d. Inner Physiology  Sharp triangular teeth in upper and lower jaws; arranged in rows that are constantly replaced  Mouth leads to pharynx with openings to gill slits  Osmoregulation accomplished by rectal gland which secretes sodium chloride; nitrogenous compounds are also retained in blood to increase solute concentrations, making more on par with seawater e. Shark Attacks  Only 32 species ( of 350) have been documented to attack humans with another 36 considered potentially dangerous; these typically are the larger size sharks; 80 % of sharks are harmless  Great white, tiger, and bull sharks are the more aggressive species  50-75 attacks occur each year, with 8-12 fatalities; in contrast 30-100 million sharks are killed every year  Attacks usually occur by sandbars, steep drop offs, or by river inlets and are associated with mistaken identity,territorial behavior, or feeding behavior 2. Rays  A. Order Rajiformes—skates, sawfish rays, electric rays, stingrays, manta rays and others Make up half of all species of Elasmobranchii b. Form and Function  Specialized for benthic life  Flattened dorsoventrally; enlarged pectoral fins are used as swimming wings  Water used in respiration enters large spiracles in head  Teeth adapted to act as rollers to crush invertebrates and sometimes small fish  Stingrays have whip-like tail with spines and venom glands  Electric rays have electric organs on sides of head C. Subclass Holocephali: Chimeras  31 species  Ratfishes  Diverged from earliest shark lineage  Mouth has flat plates for crushing invertebrates; also feeds on seaweed and small fish D. Reproduction and Development  Internal fertilization  Oviparous sharks and rays lay an egg capsule immediately after fertilization that attaches to kelp with tendrils; may take up to 2 years before mini adult hatches  Ovoviviparous sharks retain fertilized eggs in reproductive system where they are nourished by yolk of egg; “live” birth  Viviparous sharks nourish embryos with maternal bloodstream; “live” birth  Live births make it more likely more of the young survive but no other care is given after birth Embryo Development V. Superclass Osteichthyes  A. Origin, Evolution, and Diversity  Lineage developed in Silurian and now accounts for 96% of all fishes and all tetrapods  Bone replaces cartilage as fish develops  Lung or swim bladder evolved from an extension of the gut; gas filled, it aids in buoyancy  Bony operculum, a flap covering the gills that rotates outward, draws water more efficiently over them  Specialization of jaw musculature improves feeding; also unique dental characters B. Class Actinopterygii  23,600 species comprise the ray-finned fishes  Most familiar fish type a. Palaeoniscids  Earliest forms, existing from late Silurian to late Paleozoic  Small, large eyes, dorsal fin with bony rays, heterocercal tail, and interlocking scales  Survived as other fishes declined, suggesting some adaptive advantage  Gave rise to the chondrosteons and the neopterygians b. Chondrosteons  Most primitive characteristics  Heterocercal tail and ganoid scales  Living species include sturgeons, paddlefishes, and bichirs c. Neopterygians  One lineage gave rise to modern bony fishes, the teleosts  Living species are bowfin and gars which gulp air and use vascularized swim bladder to supplement the gills d. Teleosts  96 % of all living fishes; half of all vertebrates  10 mm to 17 m; up to 900 kg in weight  Found at 5,200 m to 8,000 m below sea level  Some can live in hot springs at 44 oC while others can survive in Antarctic –2 oC.  Some live in salt concentrations three times seawater; others in swamps devoid of oxygen 2. Morphological Trends Cycloid Ctenoid  Heavy armor replaced by light cycloid or ctenoid scales which made fish more mobile; some fish such as eels and catfish have completely lost scales  Fins changed to provide greater mobility and serve a variety of functions: braking, streamlining, and social communication  Homocercal tail allowed greater speed and buoyancy  Swim bladder switched from primarily respiratory to buoyancy in function  Jaw changed to increase suctioning and protrusion to secure food C. Class Sarcopterygii  1. Diversity  Only 7 species alive today; 6 lungfishes and 1 coelacanth  Early ones had lungs as well as gills, heterocercal tail; later tail became symmetrical  Skin covered in heavy scales overlaid by an enamel  Fleshy, paired lobes are used to scuttle along bottom  South American and African lungfishes can survive out of water or long periods of time 2. Coelacanth  Thought to have been extinct for 70 million years until one was dredged up off of coast of Africa in 1938  More were caught off the coast of the Comoro Islands in 1998 VI. Structural and Function Adaptations  A. Locomotion  1. Mechanism Trunk and tail muscles propel fish forward by undulations Large, rigid head minimizes yaw Very rigid body creates less yaw and a fast fish The largest fin is the tail or caudal fin for rapid forward movement. Dorsal fins on the top and anal fins underneath assist with lateral stability. Pectoral fins behind the gill covers (operculum) assist with hovering and slow turning. Pelvic fins are often small for open water swimmers but larger on bottom dwellers which use them for resting on. 2. Speed and Energy  Larger fish swim faster  Short bursts of speed are possible for a few seconds  Swimming is most economical means of motion since water buoys the animal; swimming expends 0.30 Kcal, 1.45 Kcal for walking, and 5.43 Kcal for flying B. Swim Bladder  Fish are slightly heavier than water  A shark has a very fatty liver that makes it a little buoyant; must also keep swimming to move it forward and angle itself up  Bottom dwelling fishes also lack swim bladder  Fish can control depth by adjusting volume of gas in swim bladder  Gas gland removes or adds gases from blood to remove or add gas to bladder  Some fish gulp air to fill swim bladder C. Respiration  Gill filaments are folds of tissue inside the pharyngeal cavity covered by the operculum  Continuous water flow opposite blood flow through capillaries maximizes gas exchange allowing some fish to remove 85% of O2 from H2O  Some fishes are dependent on ram ventilation as well, in which forward movement pushes more water over gills; such fish will die in an aquarium  Lungfish use lungs; eels use skin; bowfin uses gills at low temperatures and air bladder at higher temperatures; electric eel has degenerate gills and must gulp air D. Osmotic Regulation  1. Freshwater Fishes Freshwater has less salt than blood of fish so water tends to enter fish’s cells and its salts tend to leave Hyperosmotic regulators: kidney pumps out excess water and salt absorbing cells in skin remove salts from water and add to blood Euryhaline fishes live in estuary environments where they are in contact with both fresh and salt water 2. Marine Fishes  Blood has lower salt content than surrounding water so tend to lose water and gain salt  Hypo-osmotic regulators: fish drinks water bringing in more water but also salt; salt is carried by blood to gills where it is secreted by salt-secretory cells, some salt leaves in feces, and others are excreted by kidneys E. Feeding Behavior  Most time devoted to searching for food and eating  Most carnivores-feed on zooplankton, insect larvae, and other aquatic animals  Most don’t chew food since it would block flow of water across gills; swallow food whole although a few have teeth that crack prey or have some molars in throat  Some herbivores--eat plants and algae  Suspension feeders eat plankton, using gill rakers to strain food; these fish swim in large schools  Also have omnivores, scavengers, and parasites  Stomach used for storage; intestines absorb and digest nutrients F. Migration  1. Eels  Catadromous—develop in freshwater but spawn in seawater  Adult eels spawn in Sargasso Sea at depths of 300 m.  Larvae drift for 2 years before developing into elvers; males remain in brackish water; females swim hundreds of miles up rivers  Females mature for 8-15 years before returning to the sea ( 8 months to complete journey)  American eels are separate species from European eels 2. Salmon  Anadromous—living in sea but spawing in freshwater  6 Pacific salmon species, and 1 Atlantic salmon species that migrate  Pacific species migrate downstream, live in Pacific for 4 years, and then return up the same stream it was spawned in  Young fish are imprinted with the odor of their stream  Pacific salmon spawn and then die  Endangered by stream degradation, logging, pollution, and hydroelectric dams G. Reproduction  Most dioecious with external fertilization and development  Some are ovoviviparous where eggs develop in ovarian cavity—sharks, guppies, mollies  Oviparous marine fish lay large numbers of eggs, upwards of several million  Nearshore or bottom dwelling fish lay fewer, larger nonbuoyant sticky eggs  Some fish bury eggs, attach them to vegetation, incubate them in their mouths  Freshwater fish produce fewer, nonbuoyant eggs, and more care is usually provided  Many freshwater fish also have elaborate mating dances before spawning H. Growth  Egg starts to take up water after it is laid, outer layer hardens, and cell division begins  Yolk is consumed during development  Fish fry hatch carrying semitransparent yolk sac to supply food until it can forage  As fry change to adult, it may undergo dramatic changes in body shape, fins, color patterns, etc  Growth is temperature dependent; warmer fish grow more rapidly  Annual rings on scales reflect seasonal growth cycles  Most fish continue to grow throughout life and do not stop at maturity Fish Development