The vertebrates are normally divided into two groups based on the presence or ab

sence of jaws. The vast majority possess jaws and are termed Gnathostomata; jawl
ess forms are termed Agnatha. The Agnatha are represented today by the lampreys
and hagfish, organisms of little economic importance but of great interest to bi
ologists as the sole survivors of an extensive group of Paleozoic (245 to 570 mi
llion-year-old) agnathans, which includes the earliest known vertebrates. The ex
tant forms are often referred to as the cyclostomes, the fossil forms as ostraco
derms; however, these do not constitute monophyletic groups (groups including th
e common ancestor and all descendants), and some fossil Agnatha are closely rela
ted to cyclostomes.
The Physiology of Jawless Fishes
Lampreys and hagfish are similar in general appearance despite far-reaching diff
erences in internal organization. Both are elongated fish, without paired fins,
that swim with an eel-like motion and reach lengths of one meter and weights of
up to one kilogram. They possess an entirely cartilaginous skeleton; however, th
e organization of the body conforms to the basic vertebrate pattern. The lamprey
has more active life habits and, therefore, has a more developed median fin. Th
e propulsive section of the tail is longer than that of the hagfish. The lamprey s
mouth is surrounded by a suckerlike oral disc covered by small teeth on its int
ernal surface. This rasping organ allows the lamprey to attach itself to prey, o
n which it feeds by rasping away surface tissues and ingesting blood. Above and
behind the oral disc is the nasohypophysial opening, which leads into an olfacto
ry pouch and ends blindly in a dilated sac above the anterior gill pouches. Lamp
reys have seven gill pouches opening through seven gill ports. In hagfish, the m
outh is ventral and overhung by a rostrum, and it contains a dental plate that c
an be rapidly retracted, enabling the animal to bite off fragments of tissue fro
m the dead and decaying fish on which it feeds. The nasohypophysial duct passes
backward below the brain to join the pharynx, and the respiratory current passes
along it. Hagfish are quite variable in their number of gill pouches and openin
gs. Pacific hagfish have twelve pouches with separate openings (this number may
vary up to fifteen), while Atlantic hagfish have five pouches with one common op
ening. The eye is vestigial and covered by surface tissue, so it is not normally
visible.
Lampreys
There are about thirty-five species of lampreys, and all breed in freshwater and
spend the major part of their life cycle in a freshwater larval stage before tr
ansformation to an adult stage. The eggs develop into a blind larval form, the a
mmocoete, which burrows into silt banks and remains for several years filtering
microscopic particles from the water. The ammocoete is very unlike the adult, as
no suctorial disc is present; a ciliated groove on the floor of the pharynx, to
gether with a tubular endostyle, forms the feeding apparatus. After metamorphosi
s, three different types of adult lamprey are known: nonparasitic brook lampreys
; freshwater parasitic forms; and anadromous forms (living in marine conditions
but ascending rivers to spawn). The nonparasitic forms, which constitute about h
alf the known genera, have the mechanisms for parasitic feeding but do not use t
hem. The gonads mature immediately after metamorphosis, the intestine atrophies,
and the foregut remains as a solid rod. These nonparasitic forms remain dwarf a
s they cannot feed, and six to nine months after metamorphosis they reach sexual
maturity, spawn, and die. The parasitic freshwater forms live entirely in river
systems, where they feed on fish. The anadromous forms move downstream to the e
stuary or sea after metamorphosis. There, they feed voraciously and grow rapidly
, eventually returning to the river on an upstream spawning migration after one
to three years. As in all lampreys, death follows shortly after spawning. In som
e cases, it appears that populations of originally anadromous lampreys may have
become landlocked and are now entirely freshwater. That is what appears to have
happened with the lampreys (Petromyzon marinus) that now inhabit the Great Lakes
. They are capable of causing significant damage to fish populations and almost
eradicated trout and whitefish from the Great Lakes between 1950 and 1960, when

they peaked in that system, resulting in the collapse of a flourishing fishery.

Hagfish
Hagfish are purely marine fish, in contrast, and as they are benthonic (live on
the bottom) and often inhabit deep water, little is known of their reproductive
biology beyond the fact that eggs may be laid at any time of year and there appe
ars to be no larval stage. They remain buried in mud during the day, emerging at
night to feed. It appears that they normally attack only dead or dying fish, la
cking the suction apparatus that makes the lamprey so successful in its attacks
on living organisms; however, they will consume small invertebrates if they are
available. When feeding, they may show knotting behavior, during which the flexibl
e body ties into a knot in order to gain a better purchase for the tearing off o
f food fragments. One characteristic of all twenty species of hagfish is the dev
elopment of mucus pores along the body. If roughly handled or irritated, these p
ores produce copious quantities of mucus, resulting in the name slime eels for the
se animals. Neither hagfish nor lampreys are of more than local economic importa
nce.
The Fossil Record
The fossil record of hagfish and lampreys is sparse, presumably because of the l
ack of an ossified endoskeleton. Both are known only from the Pennsylvanian peri
od (280 million years ago) of Illinois and show close similarity to modern forms
. These sediments also include two other agnathans of unknown affinities, indica
ting a wide diversity of forms of this type. There is, however, an extensive rec
ord of armored agnathans (ostracoderms) fromrocks of Ordovician to Devonian age
(360 to 470 million years old). These have been divided into two main groups, th
e Osteostraci and the Heterostraci, together with some smaller groups. The Osteo
straci were a group of fish that lived in the Silurian and Devonian periods (370
to 425 million years ago) and were characterized by the presence of an armored
head-shield, the rest of the body being covered by large bony scales.The large e
yes were dorsally placed, and between them were the pineal foramen (an opening f
or a light-sensitive organ) and the nasohypophysial opening. Located laterally o
n the head-shield were sensory fields that probably were sensitive to pressure w
aves in the surrounding water. The ventrally placed mouth and dorsoventrally fla
ttened head indicate that the Osteostraci were benthonic fish, possibly feeding
by sucking organic debris or small organisms into the mouth. Paired pectoral fin
s may have acted to move the animals by rhythmic undulations. Because the head-s
hield surrounded the brain, a considerable amount of detail of brain structure a
nd cranial nerve pattern has been preserved. It has shown that the general patte
rn was very similar to that of the lamprey, implying a close relationship. The H
eterostraci were a long-ranging group whose earliest representatives occurred in
the Middle Ordovician period (470 million years ago) and represent the first kn
own record of vertebrates. Typical Heterostraci were armored over the anterior p
art of the body by variable numbers of bony plates and were further characterize
d by having only one pair of external gill openings. They were common in shallow
marine and freshwater environments during the Upper Silurian and Devonian perio
ds (360 to 425 million years ago) but became extinct at the end of the Devonian.
They seem to have been adapted to a variety of modes of life, from benthonic de
tritus feeding to cropping algae and filter feeding. The Ordovician forms are kn
own from rocks in North and South America and Australia and are united with late
r Heterostraci by the presence of the same type of acellular bone, aspidin, in t
he armor. Their exact position is uncertain, however, as they do not appear to h
ave a series of branchial (gill) openings on each side. Although it is clear tha
t the Osteostraci were closely related to modern lampreys, no relationship has b
een determined yet between hagfish and any ostracoderm group. The relationship o
f the fossil and modern agnathans to gnathostomes is also still the subject of c
onsiderable debate, though in broad terms it is accepted that lampreys are the s
ister group of gnathostomes and hagfish the sister group of lampreys and gnathos
tomes. Much further information is needed before the details of their phylogeny
can be elucidated, and continuing work on these organisms will aid understanding

of both the origin of vertebrates and the early development of major vertebrate
groups.
The Ichthyology of Jawless Fishes
Fish are generally excellent subjects for study and for the demonstration of ana
tomy, physiology, ecology, evolution, and other aspects of science. The detailed
anatomy of the cyclostomes has been known for many years, and knowledge in this
area is dependent on careful dissection of specimens. General knowledge of the
life cycle, ecology, and feeding habits of the modern forms is based on observat
ion either in the natural habitat or in aquariums. As lampreys spend most of the
ir life cycle in freshwater, understanding of their development is fairly comple
te. Hagfish, however, are marine benthonic fish that move and feed mostly at nig
ht, and hence the ability to observe them is somewhat limited and understanding
of their life cycle is incomplete. Sophisticated laboratory techniques now make
it possible to analyze the biochemistry and physiology of these organisms and co
mpare them to other chordates. Studies have also been made of their swimming met
hods, using highspeed cameras and electromyography, a technique that allows the
tracing of the electrical changes that take place in muscles when they are activ
e. Fossil Agnatha cannot be studied as completely, because only the hard parts a
re preserved in the sediments. Techniques for studying these remains have change
d very little in the past one hundred years. The bones are removed from the rock
by chipping or dissolving the surrounding sediment away. Bone is composed of ca
lcium phosphate and thus will resist some acids that can break down carbonate ro
cks. The acids most commonly used are acetic acid and formic acid, and the speci
men to be dissolved out is often backed with plastic so that it does not disinte
grate when the supporting matrix (surrounding sediment) is removed. The bone fro
m these ancient agnathans is often so well preserved that thin ground sections c
an be made and viewed through a microscope using transmitted light. The characte
rs determined by these means are used to determine relationships. Studies on the
relationships of the fossil and recent Agnatha have relied in recent years on t
he methodology termed phylogenetic systematics, or cladistics. Cladistics is disting
uished from other taxonomic methods (taxonomy is the study of interrelationships
) by the fact that it is a rigorous system in which only shared advanced charact
ers are used to show relationships. These relationships are expressed as branchi
ng diagrams termed cladograms (from the Greek klados, branch ), hence the name clad
istics. Although studies using this methodology have improved understanding of t
he relationship between modern Agnatha and the gnathostomes, the relationships o
f the fossil forms are still poorly understood.
Modern Jawless Fishes
Modern Agnatha, or cyclostomes, are relatively rare and unimportant organisms, a
lthough they are representatives of a group that was important in the early hist
ory of vertebrate evolution. For example, hagfish can be a nuisance to fishermen
, attacking and destroying bait and even the catch itself. Lampreys, because of
their active parasitic mode of life, can be a serious menace to fisheries, as ev
idenced by the sea lamprey depredations in the Great Lakes. The construction of
a canal bypassing Niagara Falls unfortunately allowed lampreys to enter the uppe
r Great Lakes from Lake Ontario, where they had been established. Once in the up
per lakes, they underwent a population explosion, probably as a result of the ab
undance of prey fishes, lack of predators, and suitability of the system for spa
wning and maintenance of larvae. The establishment of the lamprey resulted in a
serious decline of a number of fish species and the collapse of a flourishing co
mmercial fishery that has only been reversed by the establishment of control mea
sures and the use of larvicides. The fossil Agnatha, or ostracoderms, are genera
lly poorly known as a result of their incomplete preservation. Yet they do throw
some light on the earliest stages of vertebrate evolution, indicating that, as
far back as 470 million years ago, humankind s earliest ancestors were small, rath
er tadpolelike fish with an external armor of bony plates. Although they appear
in the fossil record before the earliest jawed vertebrates, or gnathostomes, it
appears that the separation into jawed and jawless forms had already occurred. M

any gaps remain, but it is to be hoped that further discoveries will enable huma
nkind to develop a clearer picture of its earliest vertebrate ancestry.