Agnathans

Hagfish

Pacific Hagfish
definitely appears to be more primitive than the lamprey
it is the most primitive living vertebrate
approximately 32 species
they are generally 1 meter or less in length
degenerate eyes covered by thick skin
but they have keen sense of smell and touch
degenerate lateral line system
lateral line detects pressured waves in water
mouth surrounded by tentacles (used for touch)
mouth has muscular tongue with keratinized teeth on it for rasping off tissue
from dead animals
form 1 to 16 gill openings on each side of the body
retains primitive kidneys (pronephros) into adulthood
(most vertebrates have this type of kidneys only during embryonic development)
conforms to osmolarity of sea water (i.e. it is isoosmotic a liquid having the
same osmotic pressure as another reference liquid)
in addition to main heart, it has several "accessory hearts" in the venous system
to help pump blood back to main heart

heart rates of all hearts are completely endogenous
no neural inputs to hearts
body has slime glands that open to outside through many pores
can secrete enormous amounts of mucous
defense mechanism
often called slime eels by fisherman
use a knotting behavior to rid themselves of the slime

Ecology

all are marine
isoosmotic (1000 mOsm body fluids)
bottom dwellers (live and feed along bottom ooze, e.g. mud)
usually in relatively deep waters (usually 300 ft or more)
scavengers
scavenge for dead or dying animals
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can use a "knotting behavior" to help it pull pieces of food off
dead prey
also uses knotting behavior to escape from predators and to rid its body of slime

Reproduction

reproductive behavior and physiology is not well understood because they live on
bottom
in relatively deep water
single long gonad
most appear to be separate sexes, but some hermaphrodites have been found
no reproductive ducts, just gonads
gametes given off into body cavity and then they exit
through a pair of genital pores into the cloaca
external fertilization (but reproduction has not been observed)
fertilized eggs appear to develop into small, but completely formed hagfish (i.e.
no larval stage)

in 1862, the Royal Danish Academy of Sciences offered a Gold Medal to
anyone who could unravel the mystery of hagfish reproduction
The award has never been claimed.
Scientifically, they represent a very interesting animal because they are
the most primitive living vertebrate

Economic Significance

meat is eaten in some part of the world
(example: Korea consumes 5 million pounds of hagfish/year)
skin is used to produce "eelskin" for leather goods
they are harvested in various areas of the world including the U.S.
certain areas of the world (e.g. Asia) have been "overfished" for hagfish

Jawless Fishes

Lamprey

Sea lamprey from Sweden
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Basic external anatomy of the lamprey

Lampreys attached to lake trout

Mouth of lamprey

approximately 41 species
maximum length of approximately 1 m
seven pairs of gills and gill openings
well-developed eyes in adult
well-developed lateral line
single heart (two chambers) with neural innervation
no accessory hearts
relatively well-developed kidneys and ion pumps on gills
body fluids maintained at approximately 300 mOsm

Ecology

Most lamprey are "anadromous".
Adults lives in marine water and then travel to freshwater to reproduce.
However, a few species are adapted completely to freshwater.
About half of the species are parasitic.
sucker-like mouth with keratinized teeth
also has tongue with teeth
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attaches to fish by suction and then uses
teeth to rasp away skin
utilizes the fish blood as food
secretes anticoagulant
some detach before fish dies

Other species of lamprey are non-parasitic.
adults only live for a few months
they metamorphose, breed, and then die
non-parasitic adults have degenerate digestive tract
(so they don't feed)

Reproduction

separate sexes with single gonad (not paired)
no reproductive ducts
marine lamprey (i.e. sea lamprey) are anadromous
migrate from marine to freshwater to breed
mate in the spring
external fertilization
egg develops into "ammocoete" larvae
(pronounced AM- mo- seat)
distinctly different from adult body form
initially thought to be a different species
and were assigned to genus Ammocetes
in many ways the larvae resemble
amphioxus and tunicate larvae
(e.g. simple chordate features)

The lancelets (subphylum Cephalochordata, and traditionally known as
the amphioxus) are a group of primitive chordates.

ammocoete larva digs into bottom of stream and filter feed long larval
stage: may feed from 3 to 17 years before they metamorphosing into
small adult lamprey
they then migrate downstream to ocean

parasitic adults may feed for about 1- 2 years
before reproducing (i.e. spawning)
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adults migrate to freshwater for spawning
die soon after spawning
some species of lamprey live entire life cycle in freshwater
example: lamprey in the Great Lakes

Economic Impact:

lamprey are eaten in some parts of the world
they have even been over-fished in some areas (e.g. certain areas in Europe)
However, in some locations, they are considered a pest, and their largest
economic impact is a negative impact on other fisheries
Example: Great Lakes Fisheries
Niagra Falls acts as a natural barrier to lamprey
between Lake Ontario and Lake Erie
During early 1900', lamprey made their way through the Great Lake system
apparently through canals built to bypass Niagara Falls.
tremendous reproductive potential: one female can produce 200,000 eggs

devastating effect on fishery
average lamprey can kill about 13 kg of fish/year
caused the collapse of Great Lake fishing industry

Lake Michigan: 1946: 5.5 million pounds ake trout captured
1953 only 402 lbs captured
Even the lamprey populations started decreasing due to lack of fish to use as
hosts.
Since the 1950's an intensive "lamprey control program" has
been conducted by Great Lakes Fishery Commission.
The control program includes

1) Currently about 50,000 to 100,000 lamprey are captured each year
females are killed
males are sterilized and released
sterile males then stimulate female to spawn, but no fertilization occurs

2) chemical larvicide is also used in streams to selectively kill ammocoete larvae

3) barriers used to block movements of lamprey
these steps have allowed fisheries to partially recover

Parasitic Lampreys
a. If marine, parasitic lampreys migrate to the sea; other species remain
in freshwater.
b. They attach to a fish by a sucker-like mouth and sharp teeth rasp
away flesh as they suck fluids.
c. They inject anticoagulant into a wound to promote flow of blood.
d. When engorged, the lamprey drops off but the wound may be fatal to
the fish.
e. Parasitic freshwater adults live 12 years before spawning and dying;
anadromous forms live 23 years.
f. Nonparasitic lampreys do not feed; their alimentary canal degenerates
as an adult, and they spawn and die.
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Sea Lamprey Invasion of the Great Lakes Region
a. No lampreys were in the U. S. Great Lakes west of Niagara Falls until
the Welland Ship Canal was built in 1829.
b. A century later, sea lampreys were first seen in Lake Erie, then
spread to all of the U.S. Great Lakes in the 1940s.
c. Lampreys preferred lake trout and destroyed this commercial species.
d. They then turned to rainbow trout, whitefish, burbot, yellow perch and
lake herring.
e. After decimating these populations, they resorted to chubs and
suckers.
f. The lamprey populations declined both from depletion of food and
from control measures.
g. Chemical larvicides were used in spawning streams; release of sterile
males is also being used.

"Gnathostomes" (i.e. jawed vertebrates)

Jaws
major development for vertebrates
Jaws are solidly braced against the cranium, but movable.
Allowed for much greater variation in behavior and adaptation
increased feeding capabilities
new food sources (more food)
increased growth, increased energy, increased activity
also allowed for digging of nests, grasping during mating, etc.

embryological and anatomical studies (including studies of fossil fishes) suggest
that jaws may have developed through modification of several sets of the anterior
gill arches
(gill arches are the support structures for the gills of fishes)

First jawed vertebrates

Fossil jawed fishes are first found in the Silurian
(approximately 420 million years ago)
In addition to jaws, early jawed fishes had
"paired fins"
greatly enhanced fishes ability to control their movements

allowed them to better use their jaws (catching prey, etc)

Jawed Fishes

two of the earliest groups of jawed fishes (both are now extinct)

a) Acanthodians
first jawed fishes in fossil record
their name refers to their stout spines
structurally they appear somewhat related to
modern groups of fishes (i.e. the Osteichthyes)
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Acanthodii (sometimes called spiny sharks) is a class of extinct fishes, having features
of both bony fish ( Osteichthyes) and cartilaginous fish (Chondrichthyes).

b) Placoderms
appear slightly later than acanthodians in fossil record
heavily armored: body covered with thick bony shields
some were large
(e.g. Dinichthys, also called Dunkleosteous: up to 10 meter long)
Both groups flourished during the Devonian, and then declined and became
extinct during the Carboniferous and Permian
The Placodermi are armoured prehistoric fishes known from fossils dating from the late
Silurian to the end of the Devonian Period. Their head and thorax were covered by
articulated armoured plates and the rest of the body was scaled or naked. Placoderms
were among the first of the jawed fish.

Placoderm

Class Chondrichthyes
sharks, skates, rays, chimaeras
fossils of early chondrichthyians date back to the Devonian Period (Age of
Fishes)
(approximately 375 million years)
Example: Cladoselache
ancient group of shark-like chondrichthyians from the Devonian
which became extinct during the Carboniferous
their structure suggested that they were fast swimmers
up to approximately 2 meters in length

General Characteristics of Living Chondrichthyes
Morphology
jawed fish with cartilaginous skeletons
paired fins and supporting skeletons
sharks have heterocercal tail
vertebral column extends into dorsal lobe of tail
skin with placoid scales (except chimaeras which lack scales)
scales with backward pointing spines
results in a tough skin that also helps reduce turbulence when
swimming
skin has mucous glands
teeth are modified placoid scales
multiple rows of teeth which are continually shed
teeth are often species or genus specific (shark can be identified
by
teeth structure)

jaws are not firmly attached to cranium (except in chimaeras)
so they can be extended when feeding
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this enlarges the size of the mouth when feeding
jaws are also powerful and can bite off pieces of prey
so sharks are not limited to prey that can be swallowed whole