DAVAO DEL NORTE STATE COLLEGE

INSTITUTE OF
AQUATIC AND
APPLIED SCIENCES

A Course Pack in
FISH 211:
ICHTHYOLOGY
Lord Martino S. Mondigo
MODULE 1
Lesson 2
Disclaimer

This module does not intend to infringe on any copyright claims and is solely for the
academic purposes of Davao del Norte State College, Its faculty, and students.

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Lesson 2

Evolution and Fish Classification

LEARNING OUTCOMES

At the end of this lesson, you should be able to:

• Illustrate the evolutionary history and taxonomic diversity of fishes

• Discuss the characteristics and features of fish belonging to different classes

INTRODUCTION

Welcome to Module No. 1 – Lesson No. 2!

Our last lesson dealt with the definition, principles, and concepts of Ichthyology
and its historical perspectives. Moreover, the lesson also tackled misconceptions about
fish and fishes. Now, we will proceed to understand the evolution and history of fishes.
Understanding fish evolution is important because of what it informs us about different
fish families, the evolution of vertebrates, and eventually, the evolution of our species.

ACTIVITY

Direction: Fish have evolved countless times throughout history, surviving mass
extinctions and significant changes to the climate and earth's structure. In this activity,
you will arrange the periods in the box below, representing evolutionary events in the
history of life on earth.

1 2 3 4 5 6 7
3

CAMBRIAN PERIOD
ORDOVICIAN PERIOD
SILURIAN PERIOD
DEVONIAN PERIOD
CARBONIFEROUS AND PERMIAN PERIOD
MESOZOIC ERA
CENOZOIC ERA

ANALYSIS

Direction: After your activity, supply the significant events in each evolutionary period.
Accomplish the table below (5 pts each):

Period Significant Events

Cambrian Period
Ordovician Period
Silurian Period

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Devonian Period
Carboniferous and Permian Period
Mesozoic Era
Cenozoic Era

ABSTRACTION

Evolution of Fish
Evolution is the process by which different living organisms are thought to have
developed and diversified from their ancestors during the earth's history. Phylogeny,
or the study of evolutionary lineages, helps scientists organize species or other groups
in ways that represent our understanding of how they evolved from common
ancestors. A common ancestor is an ancestor shared by two or more descendant
species. The evolutionary relationships among organisms are generally illustrated in a
phylogenetic tree diagram or cladogram. Evolution and diversity result from the
interactions between organisms and their environments over long periods. The
continuous adaptation of organisms to the existing diverse environments promotes a
diversity of organisms that adapted to them.
Fish began to evolve during the Cambrian explosion approximately 530 million
years ago, the earliest species classified as primitive fish initially appeared in the fossil
record. They lacked jaws and possessed rudimentary gills and spinal columns known as
notochords. Haikouichthys is one of the fish from the beginning. It possessed other
ground-breaking qualities in addition to an early spinal column. Its body was bilaterally
symmetric, meaning the left side was a mirror image of the right, and its head was
separate from its tail. Its head included a mouth and two eyes. The Agnatha, or jawless
fish, are the earliest fish lineages. During the Late Ordovician epoch, the first jawed
vertebrates most likely evolved.
In the Ordovician Period, around 480 million years ago, the spinal column began
to take on its modern form, and the first actual fish appeared in the fossil record.
Armored plates begin to develop on the head and thorax of fish. Astraspis was a jawless
fish covered with star-shaped scales from this time. Later in the Ordovician, fish first
began to develop bony jaws and, with this development, became formidable predators.
About this time, the fish line splits into two distinct lineages, the Placoderms, and
Acanthodians. The Placoderms continued the development into the bony fishes and all
other vertebrates. Acanthodians are spiny sharks. These organisms shared features of
the bony fish and sharks. In shape, their bodies resembled sharks but were covered
with bony scales that resemble the scales of modern garfish.
Late in the Silurian, the fish lines split once again as life was gaining in diversity.
The Placoderms split off the Osteichthyes, while the Acanthodians split off the
Chondrichthyes. Current bony fish are descendants of Osteichthyes, whereas modern
sharks are descendants of Chondrichthyes. The Chondrichthyes quickly diversified into
more aggressive and nimble species as the Silurian gave way to the Devonian. By the
end of the Devonian epoch, the Placoderms had vanished due to either food-related
conflict or predation by Chondrichthyes. The Devonians also saw the first of the lobe-
finned fish. These are fish similar to the Coelacanth with fleshy lobes to stiffen their fins
rather than thin bone rays like the ray-finned fish.

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a. The First Fish: Ostracoderms

The ostracoderms are the first significant vertebrate radiation well-

represented in the fossil record, other than conodonts. The earliest fossils have been
found in marine strata and belong to the Ordovician era, some 460 million years ago.
They were numerous and diverse for over 100 million years (Forey & Janvier, 1994).
There have been 600 species reported, spanning seven evolutionary lineages that
appear to be as unlike from one another as the Chondrichthyes and Osteichthyes are
now.
With a few probable exceptions, the ostracoderms were distinguished by the
absence of jaws and paired fins as well as the existence of bony armor, an internal
cartilaginous skeleton, and a heterocercal tail. Around the head, the armor was thick,
while the body armor was more like scales. The majority of ostracoderms were short
(15 cm or less in length) and dorsoventrally flattened. Their general physical makeup
suggests that they were largely bottom dwellers who survived on suckling up tiny
invertebrates and organic slime. Thelodonta, or fork-tailed thelodonts, had deep
bodies and huge eyes, indicating that free-swimming ways of living would have been
viable (Wilson & Caldwell, 1993). Ostracoderm gills, which were relatively enormous,
likely functioned as food filters and respiratory organs, similar to those of the
chordate that resembles a modern fish, amphioxus. Because most ostracoderms
lacked the control of movement provided by paired fins, they probably moved about
as much as tadpoles do. Behind the gill holes, ostracoderm did eventually acquire
bony, fin-like structures, which unquestionably improved their stability and mobility.

Figure 1. The entire body of Ostracodems

was covered in scales, like modern fish,
but the head area was more strongly
armored by the fused-together plates
into a more bony shieldc

b. The Next Fish: Placoderms

Placoderms underwent the earliest significant radiation of jawed fishes (class

Placodermi, meaning plate-skinned). Due to the weight of their hefty, bony bones are
widely documented in the fossil record. As the oldest known jawed fishes, the
Acanthodii are an important group that really made their appearance earlier in the
fossil record. They remained a relatively tiny component of the fish fauna for millions
of years. In the late Devonian period (about 350 million years ago), placoderms
controlled the oceans before going extinct totally. They were an extremely varied
group of fish, sometimes having quite odd appearances. However, based on the
design of their armor, particularly a ring of interlocking plates around the shoulder
area, they had a common ancestor (Young, 1986). Most placoderms possessed a joint-
like connection between the big bony plate on the head region and the bony plates
surrounding the trunk. Along with their characteristic layer of dermal bone plates,

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placoderms also featured paired fins, jaws, an internal body skeleton, and a
compressed dorsoventrally positioned body.

Figure 2. The placoderm fish known as Dunkleosteus, which is extinct but formerly
existed during the Late Devonian, had a maximum length of 20 feet.

c. First Chondrichthyes

The cartilaginous fishes (class Chondrichthyes) are a more manageable group

to define than the bony fishes (class Osteichthyes), partly because there are only
approximately 850 species and partly because the fossil groups (with a few
exceptions) are so poorly known that the characteristics of the living forms suffice for
the entire class. Distinctive anatomical characteristics include:

1. Cartilaginous skeleton
2. Teeth that are not fused to the jaw and that are replaced d in rows
3. Presence of unsegmented, epidermal fin rays (ceratotrichia)
4. Single, ventral nostrils on each side of the head
5. Spiral valve intestine
6. Claspers on pelvic fins of males, indicating internal fertilization.

They also lack the features of bony fish (described later in this chapter).
Fossil forms may lack some listed features; for example, well-preserved sharks from
upper Devonian deposits lack claspers. The Chondrichthyes first appeared in the
fossil record more than 400 million years ago and were common in the Devonian
period. Most of the deposits from which they are known are marine. This contrasts
with the bony fishes, which accomplished much of their early evolution in fresh
water. Within the Chondriclithyes are two distinct evolutionary lines going basic
independently as far as the Devonian: the subclass Elasmobranchii (sharks and rays)
and the subclass Holocephali (chimaeras).

Figure 3. A great white shark, an

example of the Chondrichthyes.

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c.1. Subclass Elasmobranchii: Skates and Rays

The subclass of Elasmobranchii contains skates, rays, and sharks.

Elasmobranchii members lack swim bladders and feature five to seven pairs of gill
clefts that open up to the outside. They also have rigid dorsal fins and tiny placoid
scales. The upper jaw is not fused to the skull, and the lower jaw articulates with, the
higher. The teeth are in many series. There is a tapetum lucidum in the eyes. Male
fish have grooves along each pelvic fin's inner border that serve as a clasper for sperm
transfer. These fish may be found in both tropical and arctic waters. Some are also
found in the frigid seas of the poles.

Figure 4. A common guitarfish, Rhinobatos rhinobatos, a cartilaginous fish in the family

Rhinobatidae.
c.2. Subclass Holocephali: Chaemeras

Holocephali (complete-heads) is a subclass of which the order Chimaeriformes

is the only surviving group. This group includes the rat fishes (such as the Chimaera),
rabbit fishes (an example being Hydrolagus), and elephant fishes (Callorhynchus).
Today, they preserve some features of elasmobranch life in Paleozoic times, though
in other respects, they are aberrant. They eat mollusks and other invertebrates and
are bottom-dwellers. They have a long, narrow tail and move by sweeping their big
pectoral fins. In some genera and species, the erectile spine in front of the dorsal fin
can be poisonous. The mouth is a tiny opening encircled by lips, giving the head a
parrot-like look. There is no stomach (rather, the gut is simplified, and the 'stomach'
is integrated with the intestine). The Devonian era marks the beginning of the
Holocephali fossil record. The fossil record is enormous, but the majority of them are
teeth; the body shapes of many different species are unknown or, at best, poorly
understood.

Figure 5. Chimaeras (Class Holocephali) (Photo credit: Dan Hershman)

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d. Osteichthyes – The Bony Fish

Osteichthyes, a large and diversified group of fish, have a rather extensive

fossil record. They are a challenging group to accurately characterize because of this.
Modern fonts are not distinguished by any one characteristic; rather, they are
distinguished by a typical structural pattern in conjunction with a lack of the
characteristics of the Chondrichthyes. The presence of lungs, bone, bony scales, and
lepidotrichia is one of the osteichthyan structural pattern's most recognizable
characteristics.

Figure 6. Milkfish Chanos chanos is an example of bony fish under Osteichthyes.

d.1. Sarcopterygii: Crossopterygii

The Crossopterygii subclass of bony fish, which includes the

Coelacanthiformes and Rhipidistia, is made up of both fossil and living lobe-finned or
tassel-finned fish. The former is well-known from Palaeozoic and Mesozoic rocks,
but until living examples were nabbed this century in the Indian Ocean, it was
believed they had gone extinct by the end of the Cretaceous. The Rhipidistia did go
extinct, but not before giving rise to amphibians in the Devonian. All fins of the
Crossopterygii (apart from the tail fin) are supported by moveable stalks or lobes.
Either a heterocercal or a diphycercal tail fin is present.

Figure 7. A modern coelacanth

d.2. Sarcopterygii: Dipnoi

The coelacanths (Crossopterygii) and the lungfish are the two main
groupings (super-orders) of the fleshy-finned fish subclass sarcopterygii (Dipnoi). A
single genus, Latimeria, currently represents the Crossopterygii, which first emerged
in the Devonian and were extensively dispersed throughout the Mesozoic. The

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Rhipidistia, an order of the Crossopterygii, is recognized as the ancestor of more
developed vertebrates. The Dipnoi, which now includes three species, first emerged
in the Devonian. They live in freshwater and have evolved breathing organs to take
in atmospheric air.

Figure 8. South American Lungfish

d.3. Actinopterygii: Chondrostei

The Chondrostei are the ancestors of ray-finned fishes, reaching the height of
their variety and abundance during the Carboniferous epoch. The group is more or
less a "trash can" collection of ancient fishes with numerous origins (polyphyletic),
and there is no reason to believe that it is monophyletic (Grande & Bemis, 1996).
The Neopterygii, which contains the majority of current fishes, are descended from
at least one of the old chondrostean lineages. The order Acipenseriformes, which
includes all living chondrosteans (25 species), as well as numerous extinct forms, is
represented in contemporary fauna by the two highly specialized families
Acipenseridae (sturgeons) and Polyodontidae (paddlefishes).

Figure 9. Sturgeons (above) and

paddlefishes (below) make up the
group Chondrostei.

d.4. Actinopterygii: Holostei

A class of bony fish with ray fins is called holostei. It is split into two major
clades: the Halecomorphi, which is represented by the bowfin (Amia calva), and the
Ginglymodi, which only has the gars (Lepisosteidae), which are represented by seven

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live species in two genera (Atractosteus, Lepisosteus). Over 250 million years ago,
during the Early Triassic, the clade's initial members first emerged. It was believed
that Holostei was paraphyletic. The Holostei, which both belong to the Neopterygii,
are the Teleostei's closest living relatives, according to new research. This was
discovered through Holostei's morphology, such as the existence of a paired vomer.
The chondrosteans, the other group transitional between teleosts and cartilaginous
fish, which is regarded as (at the nearest) a sister group to the Neopterigii, are more
closely related to teleosts than are the holosteans. Holostean spiracles are only
skeletal remains, and the bones have only a little ossification. The bowfin's thick
ganoid scales are more advanced than those of the gars.

Figure 10. Spotted gar, Lepisosteus oculatus

d.5. Actinopterygii: Teleostei

Teleost fish had taken over as the major species of fish in both freshwater and
ocean environments by the end of the Cretaceous period. The premaxilla and maxilla,
which make up the upper jaw's biting surface, are totally mobile in teleosts. This
mobility allows teleosts to protrude their jaws while opening their mouths.
Additionally, the totally homocercal (symmetrical) tails of teleosts set them
apart. Teleosts, which may be found from the ocean's surface to its deepest reaches,
include eels, catfish, tarpon, tuna, halibut, flounder, trout, cod, herring, salmon, and
a variety of other delectable and nutrient-rich fish.

Figure 11. Teleosts of different orders, painted by

Castelnau, 1856 (left to right, top to bottom):
Fistularia tabacaria (Syngnathiformes), Mylossoma
duriventre (Characiformes), Mesonauta acora
(Cichliformes), Corydoras splendens and
Pseudacanthicus spinosus (Siluriformes),
Acanthurus coeruleus (Acanthuriformes), Stegastes
pictus (Incertae sedis, Pomacentridae)

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APPLICATION
You are now encouraged to spare more of your time and answer the
following questions below as best as you can. In the required discussion
of your answers, it is best to cite and illustrate examples to exhaust all
areas that can support your answers (15 points each).

1. What are vertebrates? What is the oldest fish known? Provide a brief history
of fish evolution.
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
2. Give three examples of fish belonging to Super Class Agnatha and provide its
brief biology and unique features and characteristics including body structure.
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
3. Give three examples of fish belonging to Class Chondrichthyes and provide its
brief biology and unique features and characteristics including body structure.
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
4. Give three examples of fish belonging to Super Class Osteichthyes and provide
its brief biology and unique features and characteristics including body
structure.
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
5. In your own understanding, what do you think are the differences between
fish belonging to Agnatha, Chondrichthyes, and Osteichthyes?
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________

CLOSURE

Congratulations! You have successfully completed Lesson 2 of this Module. In

this lesson, you learned some evolutionary events in fish development and life on earth.
You may now proceed to the next lesson in which you are expected to learn some Fish
Structure and Form.

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REFERENCES

About: Holostei. https://dbpedia.org/page/Holostei#

Chimaeras (Class Holocephali). https://www.inaturalist.org/photos/129476

Chondrichthyes Wiki. https://chondrichthyes.fandom.com/wiki/Chondrichthyes

Common Guitarfish Rhinobatos rhinobatos (Linnaeus, 1758).

https://www.fishbase.se/FieldGuide/FieldGuideSummary.php?genusname=Rh
inobatos&speciesname=rhinobatos&c_code=132

Crossopterygii. A Dictionary of Earth Sciences Ailsa Allaby and Michael Allaby.

https://www.encyclopedia.com/plants-and-animals/zoology-and-veterinary-
medicine/zoology-general/sarcopterygians

Crossopterygian. https://www.britannica.com/animal/crossopterygian

Forey, P., and Janvier, P. (1994). Evolution of the early vertebra tes. Ame1: Scientist
82:554-565.

Helfman, G., Collette, B., Facey, D., & Bowen, B. (2009). The Diversity of Fishes: Biology,
Evolution, and Ecology. New Jersey, United State of America. Blackwell
Publishing.

History of the Earth. https://historyoftheearthcalendar.blogspot.com/2014/03/march-

15-ostracoderms.html

Introduction to the Dipnoi. https://ucmp.berkeley.edu/vertebrates/sarco/dipnoi.html

Moyle, P. & Cech, J. (2004). Fishes: An Introduction to Ichthyology (5th Edition). United
States of America. Pearson Education.

Sarcopterygii. A Dictionary of Earth Sciences Ailsa Allaby and Michael Allaby.

https://www.encyclopedia.com/plants-and-animals/zoology-and-veterinary-
medicine/zoology-general/sarcopterygians

The Evolution of Fish. https://www.fossilera.com/pages/the-evolution-of-fish

Wilson, M. H. V., and Caldwell, M. D. (1993). New Silurian and Devonian fork-tailed
"athelodonts" are jawless vertebra tes with stomachs and deep bodies. Nature
361:442-444.

Young, G. C. (1986). The relationships of placoderm fishes. Zool. J. Linn. Soc. 88:1-57.

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