Central University of Jammu

Department of Zoology
Course: Chordate biology

Origin and Evolution of Agnatha: Ostracoderms and Cyclostomes

The term Agnatha, which means jawless, was introduced in 1889 by the palaeontologist
E.D. Cope for a group that included the recent hagfishes and lampreys, and a number of
Palaeozoic, fossil fishes which, at that time, could not be proved to have possessed jaws.
Nowadays, some of these fossils are still considered as being jawless, whereas others have turned
out to be jawed vertebrates or gnathostomes. All the characters used by Cope and later scientists
to define this group are either absence of gnathostome characters (e.g. lack of jaws, lack of
horizontal semicircular canal in ear), or characters that are likely to be primitive for all
vertebrates (e.g. single median ‘nostril’). Therefore, agnathans are now merely considered as an
ensemble (group) of nongnathostome vertebrates, some of which are more closely related to the
gnathostomes than others.

In addition to the absence of jaws, modern agnathans are characterized by absence of

paired fins; the presence of a notochord both in larvae and adults; and seven or more paired
gill pouches. Lampreys have a light sensitive pineal eye (homologous to the pineal gland in
mammals). All living and most extinct Agnatha do not have an identifiable stomach.
Fertilization and development are both external. There is no parental care in the Agnatha class.
The Agnatha are ectothermic or cold blooded, with a cartilaginous skeleton, and the heart
contains 2 chambers.

General Characters:

1. Metabolism
Agnathans are ectothermic, meaning they do not regulate their own body temperature.
Agnathan metabolism is slow in cold water, and therefore they do not have to eat very much.
They have no distinct stomach, but rather a long gut, more or less homogenous throughout its
length. Lampreys feed on other fish and mammals. They rely on a row of sharp teeth to shred
their host. Anticoagulant fluids preventing blood clotting are injected into the host, causing the
host to yield more blood. Hagfish are scavengers, eating mostly dead animals. They also use a
sharp set of teeth to break down the animal. The fact that Agnathan teeth are unable to move up
and down limits their possible food types.

2. Body covering

In modern agnathans, the body is covered with skin, with neither dermal or epidermal
scales. The skin of hagfish has copious slime glands, the slime constituting their defense
mechanism. The slime can sometimes clog up enemy fishes' gills, causing them to die. In direct
contrast, many extinct agnathans sported extensive exoskeletons composed of either massive,
heavy dermal armour or small mineralized scales.

3. Appendages
Almost all agnathans, including all extant agnathans, have no paired appendages,
although most do have a dorsal or a caudal fin. Some fossil agnathans, such as osteostracans and
pituriaspids, did have paired fins, a trait inherited in their jawed descendants.

4. Reproduction
Fertilization in lampreys is external. There is no known parental care. Not much is known
about the hagfish reproductive process. It is believed that hagfish only have 30 eggs over a
lifetime. Most species are hermaphrodites. There is very little of the larval stage that
characterizes the lamprey. Lamprey are only able to reproduce once. After external fertilization,
the lamprey's cloacas remain open, allowing fungus to enter their intestines, killing them.
Lampreys reproduce in freshwater riverbeds, working in pairs to build a nest and burying their
eggs about an inch beneath the sediment. The resulting hatchlings go through four years of larval
development before becoming adults. They also have a certain unusual form of reproduction.

Origin and evolution of Agnatha

 The oldest fossil agnathans Myllokunmingia from the Lower Cambrian (530 million years
ago) of China is still considered to represent the most primitive vertebrate and is probably related
to agnathan (jawless) fish. It somewhat resembles the present-day marine hagfish (Class
Myxini). The two groups still survive today: the lampreys and the hagfish, comprising about
120 species in total. The oldest known group of vertebrates is regarded to be Agnatha which
includes the extinct ostracoderms and the living cyclostomes. Stensio (1927) believed that
cyclostomes had descended from some group of ostracoderms by the evolution of a sucking
mouth, and development of cartilage. Their structural organization has been considered higher
than that of Branchiostoma (Cephalochordata) but lower than that of Gnathostomata (jawed
vertebrates). There is increasing evidence to believe that there were two major and not so
much related groups of ostracoderms : (i) the cephalaspides having several pairs of gill slits,
pairs fin-like appendages, heterocercal tail and a single median nostril; and (ii) the pteraspids
having a single pair of gill-slits, no paired appendages, hypocercal tail and two separate nostrils.
According to Stensio (1927), the lampreys (order Petromyzontiformes) represent derivatives
of cephalaspid line and the hagfishes and slime eels (order Myxiniformes) are probably
derived from pteraspid stock.

The only fossil hagfishes and lampreys known to date are poorly preserved imprints from
the late Carboniferous (about 300 million years ago). The fossil hagfish Myxinikela (Figure
2b) shows the characteristic snout tentacles, but its body is stouter than that of living forms.
The fossil lampreys Mayomyzon (Figure 2c) and Hardistiella are much smaller than the living
ones, but are almost identical in overall shape. Living hagfishes and lampreys may thus be
regarded as ‘living fossils’.
These agnathans have been referred to by Cope as ‘ostracoderms’ (i.e. shell-like
skin), a name that still survives informally, though we now know that ostracoderms are not a
group (Figure 2d–k). Ostracoderms are the earliest known, undisputed vertebrates, and
their earliest known remains are derived from 470 million year-old Ordovician rocks.
However, ostracoderm diversity is highest during the Silurian and Devonian periods (from
430 to 365 million years). Figure 3 shows the distribution of the various ostracoderm groups
through time.
 All ostracoderms are covered with either bony plates or scales, or both, which form
the external skeleton (exoskeleton) (Figure 2d–k). Only some of them (eriptychiids, galeaspids,
osteostracans, and possibly pituriaspids) have a calcified or bony internal skeleton
(endoskeleton). In galeaspids and osteostracans, the massive, bony endoskeleton of the head
provides an accurate cast of the internal anatomy (brain, sensory capsules, nerves, blood vessels,
gills). In other groups, there is very little information on the internal structure. Apart from
osteostracans, which have true bone, with bone cells, the exoskeleton of all other ostracoderms is
devoid of bone cells and made of a hard tissue called aspidine, which is regarded as a primitive
type of bone.

Arandaspids (Arandaspida is a taxon of very early, jawless prehistoric fish which lived
during the Ordovician period. Arandaspids represent the oldest known craniates, a proposed
group of chordates that contain all chordates with a cartilage-derived skull, Figure 2f),
astraspids (Astraspida, or astraspids, are a small group of extinct armored jawless vertebrates, which
lived in the Middle Ordovician in North America), heterostracans (Heterostraci is an extinct subclass
of pteraspidomorph jawless vertebrate that lived pri3marily in marine and estuary environments, Figure
2g, h) and probably eriptychiids look like large tadpoles and are gathered into a higher group
called Pteraspidomorphi, characterized by two large, median openings are situated between two
plates on each sides.

Galeaspids, osteostracans and pituriaspids generall have a flattened, horseshoe-shaped

head, whose endoskeleton is massively ossified or calcified and covered with exoskeleton. Their
eyes are on the dorsal side of the head, and their mouth and gill openings are on the ventral side,
suggesting that they were bottom-feeders. Galeaspids (Figure 2k) have a large dorsal opening in
the anterior part of the head, which probably served for the intake of the respiratory water.

Osteostracans (Figure 2j) possessed a median dorsal ‘nostril’, which recalls the
nasohypophysial opening of lampreys and had no respiratory function. Pituriaspids (Figure 2i)
are still poorly known but they differ from galeaspids and osteostracans in lacking any
dorsal opening on top of the head.
 Anaspids (Figure 2d) are usually small, laterally compressed fishes, whose overall
aspect recalls that of lampreys. They have broad-based paired fins, yet it is unknown whether
they were mobile, like true pectoral fins. Anaspids have been regarded as possible ancestors
to lampreys because of their overall aspect, but they are poorly known and remain of debated
affinity.
Unlike all other ‘ostracoderms’, thelodonts (Figure 2e) are entirely covered with
minute scales which somewhat resemble the placoid scales of modern sharks but their internal
structure is poorly known. Thelodonts are difficult to characterize, and may possibly be an
ensemble of primitive ‘ostracoderms’, and even include ancestors of the gnathostomes.
Some of them, however, are certainly close relatives of heterostracans. Most ‘ostracoderms’
lived in shallow water, marine environments, like deltas or lagoons. Some may even have lived
in freshwater. They probably fed on minute food particles at the bottom. It is often said that
ostracoderms have been outcompeted by jawed vertebrates, but there is no clear evidence
for this, since they coexisted with large gnathostomes for about 80 million years.

Another important development was the evolution of a specialized mouth and

feeding apparatus. While the earliest agnathostomes lacked jaws, later forms developed a
number of unique structures, including circular, tooth-lined mouths and a rasping tongue.
These structures allowed agnathostomes to feed on larger and more diverse prey, including
other fish. Throughout their evolutionary history, agnathostomes have undergone a
number of adaptive radiations, diversifying into a wide range of forms and occupying a
variety of ecological niches. Today, only two groups of agnathostomes remain: hagfishes
and lampreys. These organisms continue to play important roles in marine and freshwater
ecosystems, and are the focus of ongoing research into the early evolution of vertebrates.
Fig. 2 Attempted reconstructions of fossil agnathans. (a) Haikouichthys, Lower Cambrian of
China; (b) the hagfish Myxinikela, Carboniferous of USA; (c) the lamprey Mayomyzon,
Carboniferous of USA; (d) the anaspid Rhyncholepis, Silurian of Norway; (e) the thelodont
Loganellia, Silurian of Scotland; (f) the arandaspid Sacabambaspis, Ordovician of Bolivia; (g)
the heterostracan Tartuosteus, Middle Devonian of Estonia; (h) the heterostracan Poraspis,
Lower Devonian of Spitsbergen; (i) the pituriaspid Pituriaspis, Middle Devonian of Australia; (j)
the osteostracan Zenaspis, Lower Devonian of Scotland; (k) the galeaspid Polybranchiaspis,
Lower Devonian of China; (l) the euconodont Clydagnathus, Carboniferous of Scotland. The
sizes of the fish range from about 3–4 cm (a,l) to 1 m (g) in length.
Salient features of Cyclostomata (Gr., cyklos, circular; stoma, mouth)
General Characters

1. Body elongated, eel-like.

2. Median fins with cartilaginous fin rays, but no paired appendages. Tail diphycercal.
3. Skin soft, smooth, containing unicellular mucous glands but no scales.
4. Trunk and tail muscles segmented into myotomes separated by myocommata.
5. Endoskeleton fibrous and cartilaginous. Notochord persists throughout life. Imperfect
neutral arches (arcualia) over notochord represent rudimentary vertebrae.
6. Jaws absent (group Agnatha).
7. Mouth ventral, suctorial and circular, hence the class name Cyclostomata (Gr. cyklos,
circular + stoma mouth).
8. Digestive system lacks a stomach. Intestine with a fold, typhlosole.
9. Gills 5 to 16 pairs in lateral sac-like pouches of pharynx, hence another name of class,
Marsipobranchii, Gill-slits 1 to 16 pairs.
 10. Heart 2-chambered with 1 auricle and 1 ventricle.
11. Single median olfactory sac and single median nostril. Auditory organ with 1 or 2
semicircular canals.
12. Sexes separate or united. Gonad single, large, without gonoduct.
13. Fertilization external and development is direct or with a prolonged larval stage.

Or
Although a minor element of modern marine fauna, agnathans were prominent among the
early fish in the early Paleozoic. Two types of Early Cambrian animal apparently having fins,
vertebrate musculature, and gills are known from the early Cambrian Maotianshan shales of
China: Haikouichthys and Myllokunmingia. They have been tentatively assigned to Agnatha by
Janvier. A third possible agnathid from the same region is Haikouella. A possible gnathid that
has not been formally described was reported by Simonetti from the Middle Cambrian Burgess
Shale of British Columbia. Many Ordovician, Silurian, and Devonian agnathans were armored
with heavy bony-spiky plates. The first armored agnathans-the Ostracoderms, precursors to
the bony fish and hence to the tetrapods (including humans)—are known from the middle
Ordovician, and by the Late Silurian the agnathans had reached the high point of their
evolution. Most of the ostracoderms, such as thelodonts, osteostracans, and galeaspids, were
more closely related to the gnathostomes than to the surviving agnathans, known as cyclostomes.
Cyclostomes apparently split from other agnathans before the evolution of dentine and bone,
which are present in many fossil agnathans, including conodonts. Agnathans declined in the
Devonian and never recovered.