Fishes

Chapter 24

Diversity
Fish has many usages extending beyond

what are actually considered fishes today (e.g.,

starfish, etc.).

Fishes do not form a monophyletic group.

In an evolutionary sense, can be defined as all

vertebrates that are not tetrapods.

Common ancestor of fishes is also an ancestor of
land vertebrates.

Therefore in pure cladistics, would make land vertebrates

fish.
Approximately 24,600 living species.

Adapted to live in medium 800 times denser than air.

Can adjust to the salt and water balance of their
environment.

Diversity
Evolution in an aquatic
environment both
shaped and constrained
its evolution.

Fish refers to one or

more individuals of one
species.

Fishes refers to more

than one species.

Ancestry of Fishes
Fishes have descended from an unknown freeswimming protochordate ancestor.
Agnathans including ostracoderms.
Gnathostomes derived from one group of
ostracoderms.
Four groups of gnathostomes flourished during the
Devonian, two survive today.

Fossils of Early Vertebrates

Armored, jawless
vertebrates called
ostracoderms
had defensive
plates of bone on
their skin.
One group of
ostracoderms led
to the
gnathostomes.

Fossils of Early Vertebrates

Placoderms, one group of early jawed fishes,
died out during the Carboniferous.
Left no descendents.

Fossils of Early Vertebrates

Another group, the acanthodians, were
common during the Devonian, but became
extinct during the Permian.
They were distinguished by having heavy spines on
all fins except the caudal (tail) fin.
Possible sister group of the bony fishes.

Fossils of Early Vertebrates

A third group of gnathostomes, the
cartilaginous fishes (Class Chondrichthyes)
lost the dermal armor and uses cartilage rather
than bone for the skeleton.
Sharks, skates, rays, chimaeras.

Fossils of Early Vertebrates

The last group, the bony fishes, are the
dominant fishes today.
Ray-finned fishes include most modern bony fishes.
Lobe-finned fishes contain few living species.
Includes sister group of tetrapods.
Lung fishes & coelacanths.

Origins of Bone and Teeth

Mineralization appears to have originated with
vertebrate mouthparts.

The vertebrate endoskeleton became fully

mineralized much later.

Agnathans
The least derived vertebrate lineages that still
survives are class Myxini, the hagfishes and
class Petromyzontida, the lampreys.
They lack: jaws, internal ossification, scales, and
paired fins.
Pore-like gill openings along the side of the body.

Class Myxini - Hagfish

Entirely marine.
Feeds on annelids, molluscs, crustaceans, &
dead or dying fishes.
Predators or scavengers.

Class Myxini - Hagfish

Hagfishes are jawless marine vertebrates that have a
cartilaginous skull and axial rod of cartilage derived
from the notochord.
They lack vertebrae.

Class Myxini - Hagfish

A hagfish can tie
itself in knots to
increase leverage
when burrowing into
a dead fish.

Produces large
amounts of slime.

Class Petromyzontida - Lampreys

Lampreys (Class Petromyzontida) are found
in fresh and saltwater.

Lampreys have cartilaginous segments

surrounding the notochord and arching partly
over the nerve cord.

Class Petromyzontida - Lampreys

All ascend
freshwater streams
to breed.
Marine forms are
anadromous.
Freshwater forms
move between
lakes & streams.

Class Petromyzontida - Lampreys

Lamprey larvae are called ammocoetes.
Larvae look much like amphioxus.
Possess basic chordate characteristics in simplified
form.
Suspension feeders.

Class Petromyzontida - Lampreys

Many are parasitic
as adults.
Those that are not,
do not feed as
adults.

Derived Characters of
Gnathostomes
Gnathostomes have jaws that
evolved from skeletal supports
of the pharyngeal slits.

Derived Characters of
Gnathostomes
Other characters common to gnathostomes
include:
Enhanced sensory systems, including the lateral line
system.
An extensively mineralized endoskeleton.
Paired appendages.

Fossil Gnathostomes
The earliest gnathostomes in the fossil record
are an extinct lineage of armored vertebrates
called placoderms.

Fossil Gnathostomes
Another group of jawed vertebrates called
acanthodians radiated during the Devonian
period.
Closely related to the ancestors of
osteichthyans (bony fishes).

Class Chondrichthyes
Members of class Chondrichthyes have a
skeleton that is composed primarily of
cartilage.

The cartilaginous skeleton evolved

secondarily from an ancestral mineralized
skeleton.

Subclass Elasmobranchii
The largest and most diverse subclass of

Chondrichthyes, Elasmobranchii, includes the

sharks and rays.

Subclass Elasmobranchii
Most sharks have a streamlined body and are swift
swimmers.
Heterocercal tail the upper lobe of the tail is longer
than the lower.
Placoid scales.
The upper & lower jaws have a front, functional row of
teeth and several developing rows growing behind as
replacements.

Subclass Elasmobranchii
Spiral valve in intestine slows passage of food
and increases absorptive area.

Large fatty liver aids in buoyancy.

Subclass Elasmobranchii Acute

Senses
Prey is initially detected using large olfactory organs.
Mechanorecptors in the lateral line system sense lowfrequency vibrations from far away.
Vision is important at close range.
Bioelectric fields surrounding their prey can be detected
using electroreceptors in the ampullae of Lorenzini on the
sharks head.

Subclass Elasmobranchii
All chondrichthyans have internal
fertilization.

Oviparous species lay large yolky eggs

soon after fertilization.
Some lay eggs in a capsule called a
mermaids purse that often have tendrils to
attach it to a some object.

Subclass Elasmobranchii
Ovoviviparous species retain developing
young in the uterus while they are being
nourished by the yolk.

Subclass Elasmobranchii
In viviparous species, young receive nourishment

from the maternal bloodstream through a placenta,

or from nutritional secretions produced by the
mother.
Some receive additional nutrition by eating eggs &
siblings.

Parental care ends as soon as eggs are laid or

young are born.

Subclass Elasmobranchii
Skates and rays are specialized for bottom

dwelling with a flattened body and enlarged

pectoral fins.
Gill openings on ventral surface.
Water enters through spiracles on dorsal
surface.

Subclass Elasmobranchii
Stingrays have a slender
whip-like tail with one or
more saw-edged spines
with venom glands at the
base.

Electric rays have large

electric organs that can
discharge high-amperage,
low voltage current into the
surrounding water.

Subclass Holocephali
A second subclass is composed of a few dozen
species of chimaeras, or ratfishes.
Flat plates instead of teeth.
Upper jaw fused to cranium.

Osteichthyes
Osteichthyes are the bony fishes.
Bone replaces the cartilage during development.
A swim bladder is present for controlling buoyancy
and respiration in some.
Not a monophyletic group.

Osteichthyes
Fishes breathe by drawing water over four or
five pairs of gills located in chambers covered
by a protective bony flap called the operculum.

Class Actinopterygii
Ray-finned fishes
(class
Actinopterygii)
contain all the
familiar bony fishes
more than 23,600
species.

Class Actinopterygii
The fins, supported mainly by long, flexible
rays are modified for maneuvering, defense,
and other functions.

Class Actinopterygii
Two main groups of
ray-finned fishes.
Chondrosteans
(e.g. sturgeons)
have heterocercal
tails and ganoid
scales.

Class Actinopterygii
Neopterygians
one lineage of early
neopterygians led
to the modern bony
fishes (teleosts).
Early type
neopterygians
include the bowfin
and gars.

Class Actinopterygii
The major lineage of neopterygians are teleosts,
the modern bony fishes.
Changes in fins increased maneuverability and
speed.
Symmetrical, homocercal, tail allows increased
speed.

Teleosts
Thinner, lighter cycloid and ctenoid scales
replace the heavy dermal armor of primitive
ray-finned fishes. Some (e.g. eels) lack scales.

Teleosts
Fins diversified for a
variety of functions:
camouflage,
communication,
complex
movements,
streamlining, etc.

Teleosts
The swim bladder shifted purpose from
primarily respiratory to buoyancy.
Gill arches in many diversified into pharyngeal
jaws for chewing, grinding, and crushing.

Class Sarcopterygii
Lobed-finned fishes (class Sarcopterygii)
include 2 species of coelacanths and 6 species
of lungfishes.
This group was much more abundant during the
Devonian.
Rhipidistians are an extinct group of
sarcopterygians that led to tetrapods.

Class Sarcopterygii
All early sarcopterygians had lungs as well as gills and
a heterocercal tail.
Later sarcopterygians have a continuous flexible fin
around the tail.

They have fleshy, paired lobed fins that may have been
used like legs to scuttle along the bottom.

Class Sarcopterygii
Some lungfishes can live out of the water for long
periods of time.
During long dry seasons, the African lungfish can
burrow down into the mud and secrete lots of slime
forming a hard cocoon where they will estivate until
the rains return.

Class Sarcopterygii
Coelocanths arose
during the Devonian and
peaked (max. species)
in the Mesozoic.
One genus, two species
currently.
Believed to be extinct
for 70 million years,
rediscovered in 1938.
The second species was
discovered in 1998.

Locomotion in Water
Fishes use trunk and tail musculature to propel
them through the water.

Musculature is composed of zigzag bands

called myomeres.

Locomotion in Water
Flexible fishes like eels
use a serpentine
movement.
Not very efficient for
high speed.

Fast swimmers are

less flexible.
Body undulations
limited to caudal
region.

Locomotion in Water
Many fast swimmers are streamlined with
grooves so their fins can lie flat.

Buoyancy
Sharks must move constantly to avoid sinking.
The heterocercal tail provides lift as it moves from
side to side.
Broad head and angled, stiff fins add lift.
Their large livers with fatty hydrocarbons aid in
buoyancy as well.
Liver is like a large sack of buoyant oil.

Buoyancy
Bony fishes use a gas-filled space to regulate
buoyancy the swim bladder.
Derived from a pair of lungs.
Swim bladders are absent in tunas, abyssal fishes,
many bottom dwellers.
Bony fishes will sink without the swim bladder
because they are denser than water.

Buoyancy
Fishes must be able to regulate gas inside the swim
bladder.
At depth, the gas will compress and the fish will sink.
As it rises to the surface, the gas will expand and the
fish will rise faster.

Gas may be removed in two ways.

Buoyancy
Physostomous fishes (more primitive, e.g. trout) have
a pneumatic duct that connects the swim bladder and
the esophagus.
Air can be expelled through the duct.
Gas must be secreted into the swim bladder from the
blood, although some species can gulp air to fill the
swim bladder.

Buoyancy
Physoclistous fishes
(more derived, e.g.
advanced teleosts) the
pneumatic duct has been
lost. Gas must be
absorbed by blood from
the highly vascularized
ovale.
Gas is secreted into the
swim bladder from the
blood at the gas gland.

Hearing
The bodies of fishes are

nearly the same density as

water.
Makes hearing difficult.

Weberian ossicles, found

in minnows, suckers, &
catfish, improves hearing.

Sound detection starts in

swim bladder (sound

vibrates easily in air) and
is transmitted to the inner
ear by Weberian ossicles.

Respiration
Fish gills are composed of
thin filaments covered with an
epidermal membrane that is
folded into lamellae.
Richly supplied with blood
vessels.
Located inside the pharyngeal
cavity.
Covered with an operculum
in bony fishes.
Elasmobranchs have gill slits.

Respiration
Water must be continuously pumped over the
gills.
A countercurrent system is found where the
flow of water is opposite to the flow of blood.
Deoxygenated blood encounters the freshest
water with the highest oxygen content.

Osmotic Regulation
Freshwater fishes (hyperosmotic regulators)
must have a way to get rid of water that enters
their bodies by diffusion through the gills.

Water enters the body, salts are lost by diffusion.

Water is pumped out by the opisthonephric kidney
which can form very dilute urine.
Salt absorbing cells in the gill actively move salt
from the water into the blood.

Osmotic Regulation
Saltwater fishes (hypoosmotic regulators)

have a lower blood salt concentration than the

seawater.
Tend to lose water and gain salts.
Marine teleosts drink seawater.
Salts are carried by the blood to the gills where they
are secreted out by salt-secretory cells.
Other salts are voided with feces or excreted by the
kidney.

Feeding Behavior
Most fishes are carnivores and

prey on everything from

zooplankton to large vertebrates.
Some deep-sea fishes can eat

victims twice their size an

adaptation to scarce food.
Most fishes cant chew with their
jaws (this would block water flow
over the gills), many have
pharyngeal teeth in their throats.
Large-mouthed predators can
suck prey in by suddenly opening
their mouths.

Feeding Behavior
Herbivorous fishes

eat plants and microalgae.

Most common on

coral reefs
parrotfishes,
damselfishes,
surgeonfishes.
And tropical
freshwater habitats
minnows, characins,
catfishes.

Feeding Behavior
Suspension feeders filter microorganisms from the
water using gill rakers.
Herring-like fishes are common menhaden,
herring, anchovies etc.
Many larval fishes.
Basking sharks.
Most are pelagic fishes that travel in large schools.

Feeding Behavior
Other groups are scavengers that eat dead
and dying animals,

Detritivores that consume fine particulate

organic matter,

Parasites that consume parts of other live

fishes.

Migration
Freshwater eels are catadromous, they enter
the ocean as adults, migrate to a spawning
area where they spawn & then die.
Larvae make their way back to the streams
only females enter the streams.

Migration
Anadromous salmon
spend their lives at
sea, returning to
freshwater to spawn.
Die after spawning.

Strong homing instinct

brings them to their
parent stream.
Guided by odor of
parent stream.

Reproduction
Most fishes are

dioecious with external

fertilization and
external development
oviparity.

Ovoviviparous species
(guppies, mollies,
surfperches) bear live
young after
development in the
ovarian cavity of the
female.

Reproduction
Fertilized eggs may be
pelagic and hatch into
pelagic larvae.

Large yolky benthic eggs

are often attached to
vegetation or deposited in
nests, buried, or even
carried in the mouth.
Many benthic spawners
guard their eggs.
Usually the male.

Reproduction
In some species, males defend nest sites and
perform courtship rituals to entice females to
lay their eggs in his nest. Sometimes, several
females will lay eggs in a nest.
The male will guard the eggs from predators and
will also fan them with his fins to aerate them.

Growth
Larvae may depend on the yolk sac until their
mouths and digestive systems are fully
developed.
Larvae then forage for their own food.

Growth
Larvae metamorphose into juveniles with body
shape & color patterns usually similar to the
adults.
Some species have different color patterns
in juveniles.

French Angelfishes (Pomacanthus paru) juvenile (left) and adult (right).

Growth
Growth is temperature dependent.
Fish grow faster in summer when the temperature is
warm and food is plentiful.
Growth may nearly cease during the winter.
Annual rings in scales, otoliths, and other bony
parts reflect seasonal growth.
Fish continue to grow throughout life.
Larger fishes produce more gametes.