Cuttington University

Lecture- I
Comparative
Chordate
Anatomy
Bio 212
Lecturer: Mr. Augustine S. Samorlu
0888783360/0776462002/asamorlu@[Link]
Phylogenetic relationships of Metazoans
 Introduction
• Life on Earth began some 3.6 billion years ago when a series
of reactions culminated in a molecule that could reproduce
itself.
• Although life forms may exist elsewhere in our universe or
even beyond, life as we know it occurs only on the planet
Earth.
• From this beginning have arisen all of the vast variety of living
organisms—viruses, bacteria, fungi, protozoans, plants, and
multicellular animals—that inhabit all parts of our planet.
• The diversity of life and the ability of life forms to adapt
to seemingly harsh environments is astounding.
 Comparative chordate anatomy
• Comparative chordate anatomy is a branch of biology that studies
the structures, & differences in structures of chordates.
• All forms of life are classified into five major groups known as
kingdoms.
• The generally recognized kingdoms are Monera (bacteria), Fungi,
Protista, Plantae, and Animalia.
• Within each kingdom, each group of organisms with similar
characteristics can be classified into a category known as a phylum.
• The phylum Chordata is the only group of multicellular animals that
possess the following combination of structures: (1) a dorsal hollow
nerve cord; (2) a flexible supportive rod (notochord) running
longitudinally through the dorsum just ventral to the nerve cord; (3)
pharyngeal slits or pharyngeal pouches; and (4) a postanal tail.
General Summary of
HEMICHORDATA
AND CHORDATA
 Phylum Hemichordata (hemi-kor-datah)
• They are widely distributed in shallow, marine,
tropical waters and deep, cold waters; soft
bodied and wormlike; epidermal nervous
system; most with pharyngeal slits.
• Class Enteropneusta (enter-op-nustah)
Shallow-water, wormlike animals; inhabit
burrows on sandy shorelines; body divided into
three regions: proboscis, collar, and trunk.
• Acorn worms (Balanoglossus, Saccoglossus).
• About 70 species.
• Class Pterobranchia (tero-brangke-ah) With or
without pharyngeal slits; two or more arms;
often colonial, living in an externally secreted
encasement. Rhabdopleura. About 20 species.
• Class Planctosphaeroidea
(plankto-sfer-roide-ah) Spherical body with
ciliary bands covering the surface; U-shaped
digestive tract; coelom poorly developed;
planktonic.
• Only one species is known to exist
(Planctosphaera pelagica).
 Phylum Chordata (kor-datah) (L. chorda, cord)
• Occupy a wide variety of marine, freshwater, and
terrestrial habitats.
• A notochord, pharyngeal slits, a dorsal tubular nerve
cord, and a postanal tail are all present at some time in
chordate life histories.
• About 45,000 species.
• Subphylum Urochordata (uro-kor-datah)
• Notochord, nerve cord, and postanal tail present only in
free swimming larvae; adults sessile, or occasionally
planktonic, and enclosed in a tunic that contains some
cellulose; marine. Sea squirts or tunicates.
•Class Ascidiacea (as-ide-ase-ah) All
sessile as adults; solitary or colonial;
colony members interconnected by
stolons.
•Class Appendicularia
(a-pendi-ku-lare-ah) (Larvacea)
(lar-vase-ah)
•Planktonic; adults retain tail and
notochord; lack a cellulose tunic;
epithelium secretes a gelatinous covering
•Class Sorberacea (sorber-ase-ah)
•Ascidian-like urochordates possessing
dorsal nerve cords as adults; deep
water; carnivorous. Octacnemus.
•Class Thaliacea (tale-ase-ah)
•Planktonic; adults are tailless and barrel
shaped; oral and atrial openings are at
opposite ends of the tunicate; muscular
contractions of the body wall produce
water currents.
 Subphylum Cephalochordata (sefa-lo-kor-datah)
• Body laterally compressed and transparent; fishlike; all
four chordate characteristics persist throughout life.
Amphioxus (Branchiostoma).
• About 45 species.
• Subphylum Vertebrata (verte-bratah)
• Notochord, nerve cord, postanal tail, and pharyngeal
slits present at least in embryonic stages; vertebrae
surround nerve cord and serve as primary axial
support; skeleton modified anteriorly into a skull for
protection of the brain.
• Class Cephalaspidomorphi
(sefah-laspe-do-morfe)
• Fishlike; jawless; no paired appendages;
cartilaginous skeleton; sucking mouth with teeth
and rasping tongue. Lampreys.
• Class Myxini (miks˘ı-ne)
• Fishlike; jawless; no paired appendages; mouth
with four pairs of tentacles; olfactory sacs open
to mouth cavity; 5 to 15 pairs of pharyngeal slits.
Hagfishes.
 Class Chondrichthyes (kon-drikthi-es)
• Fishlike; jawed; paired appendages and cartilaginous
skeleton; no swim bladder. Skates, rays, sharks.
• Class Osteichthyes (oste-ikthe-es)
• Bony skeleton; swim bladder and operculum present.
Bony fishes.
• Class Amphibia (am-fibe-ah)
• Skin with mucoid secretions; possess lungs and/or
gills; moist skin serves as respiratory organ; aquatic
developmental stages usually followed by
metamorphosis to an adult. Frogs, toads,
salamanders.
• Class Reptilia (rep-tile-ah)
• Dry skin with epidermal scales; amniotic eggs;
terrestrial embryonic development. Snakes, lizards,
alligators.
• Class Aves (avez)
• Scales modified into feathers for flight; efficiently
regulate body temperature (endothermic); amniotic
eggs. Birds.
• Class Mammalia (mah-male-ah)
• Bodies at least partially covered by hair; endothermic;
young nursed from mammary glands; amniotic eggs.
Mammals.
 Phylum Hemichordata
• The Phylum Hemichordata lacks two major
features that distinguish them from other
chordates, namely; a post-anal tail and
notochord, thus, classifying them as
half-chordates.
• The body of hemichordates are divided into
three parts: Trunk, Collar and Proboscis
Hemichordata
 Characteristics
• Bilaterally symmetrical and triploblastic
• Their body wall consists of single-layer epidermis
• True body cavity
• The proboscis has a glomerulus, which is the excretory organ.
• Have a complete digestive tract.
• Have an open system of circulation.
• Excretion takes place in the proboscis by a single glomerulus.
• Dioceous or hermaphrodite.
• Sexual reproduction.
• One to several pairs of Gonads.
• Respiration occurs through their gills.
• Free swimming tornaria larva with external fertilization.
• Dorsal, sometimes tubular, nerve cord
 Classes
• Subphylum Hemichordata is divided into three classes;
• Enteropneusta : The class Enteropneusta consist of
acorn worms
• Pterobranchia: The class Pterobranchia may include the
graptolites
• Planctosphaeroidea: This class includes only the
species planctosphaeroidea pelagica
 Major divisions of chordates
• The phylum chordata is divided into three major
subphyla: Urochordata, Cephalochordata, and
Vertebrata.
• The Urochordata and Cephalochordata consist of
small, nonvertebrate marine animals and are
often referred to collectively as protochordates.
• To clearly understand and compare their
evolutionary significance in relation to the
vertebrates, it is necessary to briefly discuss their
characteristics.
 Urochordata (tunicates):
• Adult tunicates, also known as sea squirts, are
mostly sessile, filter-feeding marine animals whose
gill slits function in both gas exchange and feeding
.
• Water is taken in through an incurrent siphon,
goes into a chamber known as the pharynx, and
then filters through slits into the surrounding
atrium.
• Larval tunicates, which are free-swimming, possess
a muscular larval tail that is used for propulsion.
• The name Urochordate is
derived from the Greek
oura, meaning tail, and
the Latin chorda, meaning
cord; thus, the
“tail-chordates.”
• When the larva
transforms into an adult,
the tail, along with its
accompanying notochord
and most of the nerve
cord, is reabsorbed
 Cephalochordata (lancelet; amphioxus)
• Cephalochordates are small (usually less than 5 cm long),
fusiform (torpedo-shaped) marine organisms that spend
most of their time buried in sand in shallow water.
• Their bodies are oriented vertically with the tail in the sand
and the anterior end exposed.
• A well-developed notochord and long dorsal hollow nerve
cord extend from the head (cephalo) to the tail and are
retained throughout life.
• The numerous pharyngeal gill slits are used for both
respiration and filter feeding.
• Cephalochordates have a superficial resemblance to the
larvae of lampreys (ammocoete), which are true vertebrates.
• Serially arranged blocks of muscle known as
myomeres occur along both sides of the body of the
lancelet.
• Because the notochord is flexible, alternate
contraction and relaxation of the myomeres bend the
body and propel it.
• Other similarities to vertebrates include a closed
cardiovascular system with a two-chambered heart,
similar muscle proteins, and the organization of cranial
and spinal nerves.
• No other group of living animals shows closer
structural and developmental affinities with
vertebrates.
• However, even though cephalochordates now are
believed to be the closest living relatives of
vertebrates, there are some fundamental
differences.
• For example, the functioning units of the excretory
system in cephalochordates are known as
protonephridia.
• They represent a primitive type of kidney design that
removes wastes from the coelom.
• In contrast, the functional units of vertebrate kidneys,
which are known as nephrons, are designed to
remove wastes by filtering the blood.
 SubphylumVertebrata (vertebrates)
• Vertebrates are chordates with a “backbone”—either a
persistent notochord as in lampreys and hagfishes, or a
vertebral column of cartilaginous or bony vertebrae that
more or less replaces the notochord as the main support
of the long axis of the body.
• All vertebrates possess a cranium, or braincase, of
cartilage or bone, or both.
• The cranium supports and protects the brain and major
special sense organs.
• Many authorities prefer the term Craniata instead of
Vertebrata, because it recognizes that hagfish and
lampreys have a cranium but no vertebrae.
• In addition, all vertebrate embryos pass through a
stage when pharyngeal pouches are present.
• Most living forms of vertebrates also possess paired
appendages and limb girdles.
• Vertebrate classification is ever-changing as
relationships among organisms are continually being
clarified.
• For example, hagfish and lampreys, which were
formerly classified together, each have numerous
unique characters that are not present in the other.
• They have probably been evolving independently for
many millions of years.
• Wide-ranging and diverse vertebrates successfully
inhabit areas from the Arctic (e.g., polar bears) to the
Antarctic (e.g., penguins).
• Many vertebrates are aquatic (living in salt water or
fresh water); others are terrestrial (living in forests,
grasslands, deserts, or tundra).
• Some forms, such as blind salamanders
(Typhlomolge, Typhlotriton, Haideotriton), mole
salamanders (Ambystoma), caecilians
(Gymnophiona), and moles (Talpidae) live beneath
the surface of the Earth and spend most or all of their
lives in burrows or caves.
 Body temperature
• Fishes, salamanders, caecilians, frogs, turtles, and snakes are
unable to maintain a constant body temperature independent
of their surrounding environmental temperature.
• Thus, they have a variable body temperature, a condition
known as poikilothermy, derived from heat acquired from the
environment, a situation called ectothermy.
• Although lizards are poikilothermic, many species are very
good thermoregulators.
• Birds and mammals, on the other hand, are able to maintain
relatively high and relatively constant body temperatures, a
condition known as homeothermy, using heat derived from
their own oxidative metabolism, a situation called
endothermy.
Chordates characters
 Vertebrate Evolution
• Biologists attempt to classify living things according to their
evolutionary relationships.
• The first step in classification is the grouping together of related forms;
the second is the application of names to the groups.
• Some refer to the first step as systematics and the second as taxonomy;
others use the two terms interchangeably to describe the entire
process of classification.
• Systematics comes from the Latinized Greek word systema, which was
applied to early systems of classification.
• It is the development of classification schemes in which related kinds of
animals are grouped together and separated from less-related kinds.
• Taxonomy is derived from two Greek words: taxis, meaning
“arrangement,” and homos, meaning “law.”
• It is the branch of biology concerned with applying names to each of
the different kinds of organisms.
 BINOMIAL NOMENCLATURE
• In 1758, Carl von Linne, (Carolus Linnaeus) published the
tenth edition of a book on animals entitled Systema
Naturae.
• In that edition, the binomial system of nomenclature
(two names) was applied consistently for the first time.
• The scientific name (binomen) of every species consisted
of two Latin or Latinized words: The first was the name of
the genus to which the organism was assigned, and the
second was the trivial name.
• He grouped all the classes of animals together as
members of the animal kingdom, as distinct from the
plant kingdom.
 • The classes established by Linnaeus were as follows:
I. Quadrupeds Hairy body; four feet; females viviparous,
milk-producing
II. Birds Feathered body; two wings; two feet; bony beak; females
oviparous
III. Amphibia Body naked or scaly; no molar teeth; other teeth always
present; no feathers
IV. Fishes Body footless; possessing real fins; naked or scaly
V. Insects Body covered with bony shell instead of skin; head
equipped with antennae
VI. Worms Body muscles attached at a single point to a quasi-solid
base
• Classes I, II, and IV correspond to the traditional evolutionary
taxonomic classes (mammals, birds, and fishes) used today.
• Class III, however, included both amphibians and reptiles.
 CLASSIFICATION
• The basic unit of classification, and the most important
taxonomic category, is the species.
• Related species are grouped together in a genus.
• A genus, therefore, is a group of closely related species or a
group of species that have descended from a common
ancestral group (or species).
• For example, the domestic dog together with wolves and
jackals make up the genus Canis.
• When referring to the dog, the trivial name is added—Canis
familiaris; the wolf, a close relative, is Canis lupus.
• The name of a species is always a binomen and consists of the
genus and the trivial name.
• The sequence from top to bottom indicates decreasing
scope or inclusiveness of the various levels.
• For example:
Kingdom — Animalia
Phylum — Chordata
Class — Mammalia
Order — primate
Family — Hominid
Genus — Homo
species — Sapiens
• These seven categories are considered essential to
defining the relationships of a given organism.
The evolution
of the major
groups of
hagfishes,
lampreys
fishes and
their
relationships
to each other,
and to the
amphibians
 Fish
Do a well labelled diagram of the
internal anatomy of a fish
 Cyclostomes/Agnathans
• Ostracoderms are extinct agnathans that belonged to
several classes.
• The fossils of predatory water scorpions (phylum
Arthropoda) are often found with the fossil of ostracoderms.
• Ostracoderms were bottom dwellers, often about 15 cm
long.
• Most were probably filter feeders, either filtering suspended
organic matter from the water or extracting annelids and
other animals from muddy sediments.
• Some ostracoderms may have used bony plates around the
mouth in a jawlike fashion to crack gastropod shells or the
exoskeletons of arthropods.
 Class Myxini
• Hagfishes are members of the class Myxini
• They live buried in the sand and mud of marine
environments, where they feed on soft-bodied
invertebrates and scavenge dead and dying fish .
• When hagfishes find a suitable fish, they enter the fish
through the mouth
and eat the contents of the body,
leaving only a sack of skin and bones.
• They are known to have excessively slimy bodies.
• Most zoologists now consider the hagfishes to be the
most primitive group of vertebrates.
 Class Cephalaspidomorphi
• Lampreys are agnathans in the class Cephalaspidomorphi
• They are common inhabitants of marine and freshwater environments in
temperate regions.
• Most adult lampreys prey on other fishes, and the larvae are filter feeders.
• The mouth of an adult is suckerlike and surrounded by lips that have
sensory and attachment functions.
• Lampreys have salivary glands with anticoagulant secretions and feed
mainly on the blood of their prey.
• Some lampreys, however, are not predatory.
• Members of the genus Lampetra are called brook lampreys.
• The larval stages of brook lampreys last for about three years, and the
adults neither feed nor leave their stream.
• They reproduce soon after metamorphosis and then die.
• Adult sea lampreys live in the ocean or the Great Lakes.
 Gnathostome Fishes
• The two groups of living gnathostome (jawed)
fishes are the Chondrichthyes or cartilaginous
fishes (sharks, skates, rays, and ratfishes), and
the Osteichthyes or bony fishes.
• Fishes are the most diverse group of
vertebrates, with approximately 26,000 species
of bony and cartilaginous fishes extant in the
world today.
Gnathostome Fishes
 Class Chondrichthyes
• Members of the class Chondrichthyes (Gr. chondros, cartilage
ichthyos, fish) include the sharks, skates, rays, and ratfishes .
• Most chondrichthians are carnivores or scavengers, and marine.
• In addition to their biting mouthparts and paired appendages,
chondrichthians possess epidermal placoid scales and a
cartilaginous endoskeleton.
• The subclass Elasmobranchii (Gr. elasmos,plate metal branchia,
gills), which includes the sharks, skates, and rays, has about seven
hundred species.
• Tough skin with dermal, placoid scales covers sharks.
• These scales project posteriorly and give the skin a tough,
sandpaper texture.
• (In fact, dried shark skin has been used for sandpaper.)
reef shark AND stingray
• Crowns of sharks` teeth in different species may be
adapted for shearing prey or for crushing the shells of
molluscs.
• Sharks range in size from less than 1 m (e.g., Squalus, the
laboratory dissection specimen) to greater than 10 m
(e.g., basking sharks and whale sharks).
• The largest sharks are not predatory but are filter
feeders.
• They have pharyngeal-arch modifications that strain
plankton.
• The fiercest and most feared sharks are the great white
shark (Carcharodon) and the mako (Isurus).
• Skates and rays are specialized for life on the ocean floor
using their blunt teeth to feed on invertebrates.
• Their most obvious modification for life on the ocean floor is
a lateral expansion of the pectoral fins into winglike
appendages.
• Locomotion results from dorsoventral muscular waves that
pass posteriorly along the fins.
• Frequently, elaborate color patterns on the dorsal surface of
these animals provide effective camouflage.
• The sting ray (Dasyatis) has a tail modified into a defensive
lash; the dorsal fin persists as a venomous spine.
• Also included in this group are the electric rays (Narcine and
Torpedo) and manta rays (Manta).
 Subclass Holocephali
• A second major group of chondrichthians, in the subclass
Holocephali (Gr. holos, whole kephalidos, head), contains about
30 species.
• A frequently studied example, Chimaera, has a large head with a
small mouth surrounded by large lips.
• A narrow, tapering tail has resulted in the common name “ratfish.”
• Holocephalans diverged from other
chondrichthians nearly 300 million years ago.
• Since that time, specializations not found in other elasmobranchs
have evolved, including a gill cover, called an operculum, and teeth
modified into large plates for crushing the shells of molluscs.
• Holocephalans lack scales.
Subclass Holocephali
 Class Osteichthyes
• Members of the class Osteichthyes (Gr. osteon, bone
ichthyos, fish) are characterized by having at least some
bone in their skeleton and/or scales, bony operculum
covering the gill openings, and lungs or a swim bladder.
• Members of the subclass Sarcopterygii (Gr. sark, flesh
pteryx, fin) have muscular lobes associated with their fins
and usually use lungs in gas exchange.
• One group of sarcopterygians are the lungfishes
• When freshwater lakes and rivers begin to stagnate and
dry, these fishes use lungs to breathe air
lungfish
 The coelacanths
• A second group of sarcopterygians are the
coelacanths.
• The discovery of this fish, Latimeria chalumnae, was
a
milestone event because Latimeria is probably
the closest living fish relative of terrestrial vertebrates.
• It is large—up to 80 kg—and has heavy scales.
• Ancient coelacanths lived in freshwater lakes and
rivers.
Latimeria
 subclass Actinopterygii
• The subclass Actinopterygii (Gr. aktis,ray pteryx, fin)
contains fishes that are sometimes called the rayfinned
fishes because their fins lack muscular lobes.
• They usually possess swim bladders; gas-filled sacs along
the dorsal wall of the body cavity that regulate buoyancy.
• Ancestral chondrichthians had a bony skeleton, but living
members, the sturgeons and paddlefishes, have
cartilaginous skeletons.
• Sturgeons feed on invertebrates that they stir up from
the sea or riverbed using their snouts.
Sturgeons And paddlefish
 CIRCULATION
• All vertebrates have a closed circulatory system in which
a heart pumps blood, with red blood cells containing
hemoglobin, through a series of arteries, capillaries, and
veins.
• The vertebrate heart develops from four embryological
enlargements of a ventral aorta.
• Fishes have a closed circulatory system with a heart that
pumps blood around the body in a single loop-from the
heart to the gills, from the gills to the rest of the body,
and then back to the heart. The fishes heart consist of
four parts: the sinus venosus, atrium, ventricle, and the
bulbus arteriosus.
 Gas Exchange
• Fishes live in an environment that contains less than 2.5% of
the oxygen present in air.
• To maintain adequate levels of oxygen in their bloodstream,
fishes must pass large quantities of water across gill surfaces
and extract the small amount of oxygen present in the water.
• Gill filaments extend from each gill arch and include vascular
folds of epithelium, called pharyngeal lamellae.
• Branchial arteries carry blood to the gills and into gill
filaments.
• The arteries break into capillary beds in pharyngeal lamellae.
• Gas exchange occurs as blood and water move in opposite
directions on either side of the lamellar epithelium.
NUTRITION AND THE DIGESTIVE SYSTEM
• Most modern fishes are predators and spend much of their life
searching for food.
• Some fishes feed on invertebrate animals floating or swimming
in the plankton or living in or on the substrate.
• To capture prey, fishes often use the suction that closing the
opercula and rapidly opening the mouth creates, which
develops a negative pressure that sweeps water and prey
inside the mouth.
• An enlargement, called the stomach, stores large, often
infrequent, meals.
• The small intestine like others vertebrates, however, is the
primary site for enzyme secretion and food digestion.
 Swim Bladders and Lungs
• Swim bladders are more or less oval, soft-walled sacs in the
abdominal cavity just below the spinal column.
• They may be paired or unpaired and may be partitioned into
chambers.
• A pneumatic duct, when present
in adults, usually connects the swim
bladder to the esophagus; it may also,
connect the swim bladder to the pharynx or even to the
stomach.
• Fishes in which their swim bladder are connected to the
digestive tract via a pneumatic duct are termed physostomes
and include many of the more ancestral soft-rayed teleosts
 Buoyancy Regulation
• Fishes maintain their vertical position in a column of
water in one or more of four ways;
• Incorporate low-density compounds into their tissues.
Use fins to provide lift
• The pectoral fins of a shark are planning devices that help
to create lift as the shark moves through the water.
• The reduction of heavy tissues in fishes.
• The bones of fishes are generally less dense than those of
terrestrial vertebrates.
The swim bladder
• A fish regulates buoyancy by precisely controlling the
volume of gas in its swim bladder.
 NERVOUS AND SENSORY FUNCTIONS
• The central nervous system of fishes, as in other vertebrates,
consists of a brain and a spinal cord.
• Sensory receptors are widely distributed over the body.
• In addition to generally distributed receptors for touch and
temperature, fishes possess specialized receptors for olfaction,
vision, hearing, equilibrium and balance, and for detecting water
movements.
• Openings, called external nares, in the snouts of fishes lead to
olfactory receptors.
• Recent research has revealed that some fishes rely heavily on their
sense of smell.
• For example, salmon and lampreys return to spawn in the streams
in which they hatched years earlier.
• The eyes of fishes are similar in most aspects of structure to those
in other vertebrates.
• They are lidless with round lenses.
• Focusing requires moving the lens forward or backward in the eye.
• Fishes lack the outer and/or middle ear, which conducts sound
waves to the inner ear in other vertebrates.
• Running along each side and branching over the head of most
fishes is a lateral-line system.
• The lateral-line system consists of sensory pits in the epidermis of
the skin that connect to canals that run just below the epidermis.
• Lateral lines are used to detect either water currents, a predator
or a prey that may be causing water movements in the vicinity of
the fish.
• Fishes may also detect low-frequency sounds with these
 EXCRETION AND OSMOREGULATION
• Fishes, like all animals, must maintain a proper balance of
electrolytes (ions) and water in their tissues.
• This osmoregulation is a major function of the kidneys and gills of
fishes.
• Kidneys are located near the midline of the body, just dorsal to the
peritoneal membrane that lines the body cavity.
• As with all vertebrates, the excretory structures in the kidneys are
called nephrons.
• Nephrons filter blood borne nitrogenous wastes, ions, water, and
small organic compounds across a network of capillaries called a
glomerulus.
• The filtrate then passes into a tubule system, where essential
components may be reabsorbed into the blood.
• The filtrate remaining in the tubule system is then excreted.
• Freshwater fishes live in an environment containing
few dissolved substances.
• Osmotic uptake of water across gill, oral, and
intestinal surfaces and the loss of essential ions by
excretion and defecation are constant.
• To control excess water buildup and ion loss,
freshwater fishes never drink and only take in water
when feeding.
• Reabsorption of some ions and organic compounds
follows filtration.
• Ions are still lost, however, through the urine and by
diffusion across gill and oral surfaces.
 REPRODUCTION AND DEVELOPMENT
• The onset of reproductive activity (generation time) in fishes
varies with sex.
• In many species, especially those with sexual differences in size
(e.g., species of sharks), males mature at a smaller size and
younger age than do females.
• Various means of sexual recognition have evolved among
fishes.
• Some, such as female stingrays (Dasyatis), passively emit
electric fields that attract males .
• A difference in form or color between the sexes of a given
species is known as sexual dimorphism
• This sexual dimorphism is particularly evident in those fishes
that bear live young, such as members of the mosquitofish.
• In others, especially species in which males compete for
females, the males are the larger sex (e.g., many coral-reef
• Most fishes do not establish long-lasting pair bonds.
• Males and females come together to breed and then go their
separate ways.
• Some female mosquitofish (Gambusia) will mate with dozens of
males during one breeding cycle.
• Offspring of these multiple unions are genetically more diverse
than offspring of females that mate with just one male.
• Females initially lay a small quantity of eggs at a male-guarded
site.
• If the male guards the eggs successfully, she will continue laying
at his site.
• Most female bony fishes release eggs into the water, where
sperm are deposited by males.
• Sperm exhibit much variation in shape.
• Fertilization in fishes is usually external.
• As females release eggs, sperm are released by one or
more males in the same general area.
• In some teleosts, such as Fundulus, the male and female
interlock their anal and pelvic fins and then execute
external fertilization.
• Internal fertilization occurs in most cartilaginous fishes
and in some teleost fishes.
• Males of some teleost species possess an intromittent
organ known as a gonopodium.
• Gonopodia have grooved passageways that guide sperm
into the female during copulation.
• Sharks and rays have modified pelvic fins known as
claspers
• A clasper is inserted into the female, and sperm are
transported along a groove into her cloaca.
• Many fishes with internal fertilization, such as guppies
(Poecilia reticulata), dogfish sharks (Squalus), and
coelacanths (Latimeria), carry the embryos until the time
of birth.
• Some fishes are hermaphroditic, a condition in which one individual is
both male and female.
• Some hermaphrodites possess both male and female sex organs at
the same time and are called synchronous hermaphrodites; others
change sex as they grow and are called sequential hermaphrodites.
• Fertilization is external, and each member of the pair assumes a
specific sex role associated with the release of eggs or sperm.
• Individuals release only one type of gamete during a spawning
episode.
• Individuals of other species, such as black hamlets (Hypoplecturus
nigricans), take turns releasing eggs for their partners to fertilize, a
mating system referred to as “serial monogamy.”
• The most common form of sequential hermaphroditism, called
protogyny, occurs when females turn into males.
• A less common form is protandry, a change from male to female.
 Prenatal Development
Oviparous Species
• All skates, some sharks, and many bony fishes are oviparous.
• Some female sharks that can store sperm release fertilized
eggs over an extended period of time.
Viviparous Species
• Viviparity is defined as the retention of fertilized eggs within
the female reproductive tract.
• Embryos may be enclosed in a thin membrane or envelope
along with a yolk supply, or they may be nourished by direct
transfer of nutrients from the mother.
• Young are born alive.
• Most sharks, all rays, sawfish, guitarfish, and some bony
fishes, including the coelacanth are viviparous.
 Duration of Embryonic Development
• Live-bearing sharks and rays have gestation periods of 6 to 22
months.
• Hatching in oviparous fishes, however, may be as short as 1–2 days in
gouramis and coral fishes or as long as 16 months in some sharks and
rays.
Hatching and Birth
• Most fish embryos produce hatching enzymes that assist in digesting
the membrane enclosing them at the time of hatching.
• In many bony fishes, secretions come from special glands on the head
or inside the mouth.
• The sex of some fishes, such as the Atlantic silverside, is determined
in part by the temperature of the water when the young are born.
• Low temperatures early in the season produce females; higher
 Growth and Metamorphosis
• The life of an oviparous fish is divided into four stages: embryo,
larva, juvenile, and adult.
• Most of the embryonic stage is passed within the egg, although it
may continue for a short time after hatching while the embryo’s
nutrition is still derived from yolk.
• The larval period begins when the fish is able to catch food
organisms.
• During this time, the skeleton and other organ systems become
fully formed, and the fins develop.
• In the juvenile stage, the fish and its organ systems grow
considerably in size; a period of rapid growth.
• During this stage, which lasts until the gonads mature, the fishes
may be distinctively colored.