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MARINE
ICHTHYOLOGY
A Laboratory Manual
Dr Fadwa Abou-Seedo
Dr M N V Subrahmanyam
Department of Biological Sciences
Faculty of Science
Kuwait University
2013
Preface
Course Objectives: This course provides a general overview of the biology of fishes,
including aspects of their taxonomy, anatomy, physiology, behavior, natural history,
ecology, management, and conservation. Laboratory and field studies complement
and expand on concepts covered in lecture, and will expose you to a wide variety of
techniques used to study fishes. When you complete this course, you should (1) have
a broad understanding of the biology of fishes; (2) be able to identify major
taxonomic groups of fishes; (3) be able to identify most local fishes; and (4)
understand how to apply a broad array of field and laboratory techniques to the study
of fishes.
Course Overview: This course will combine a series of lectures with extensive
hands-on laboratory and field work to teach you about fishes. Our intention is to
involve you as an active participant – in lectures and in field and lab studies – because
doing something is the best way of learning it. The three portions of the course
(lecture, lab, and field) are integrated and you must take them all in the semester.
In lectures you will learn fundamental concepts in fish biology. You are
encouraged and expected to ask questions during lecture. We will cover material in
your textbook, assigned research papers, and outside material. You will learn how to
do research on fishes with a series of laboratory and field research projects. You will
learn methods for studying ichthyology in the lab and field; you will learn how to
analyze and present the data you have collected; and you will learn how to write clear,
effective, scientific research reports.
Lab practical exams: There will be three lab practicals exams. For these you will be
required to identify local species, as well as important families and higher groupings
(e.g., orders), and anatomy (e.g., fins, scales, organs, and bones). You will be required
to identify both dead and live fishes. Because of the substantial amount of time
required to set up lab practicals, there will be no make-ups.
II
Reports: You will be required to write 2 papers in the form of published scientific
papers. One of these papers (the lab report) will be based on a field study that we
conduct together as a class. The second paper, the research report, will be an
independent effort on your part. You will have the choice of conducting an
independent research project (in the field or lab) or an extensive, synthetic literature
review on a topic of your choice. In either case, when you have completed your
research project, in addition to writing the paper, you will give a 10-minute oral
presentation of your findings (as one would at a scientific meeting). We will discuss
the format and expectations for these papers and presentation in more detail later in
the semester.
Field Trips:
There will be field trips during the semester and they are all mandatory. Most of these
trips will extend beyond normal class hours and one is on a Saturday. You will not be
able to make up missed field trips. The field trips are designed to teach you how to
study fishes in nature and are essential for your understanding and appreciation of the
subject matter. Come prepared for field trips. Be prepared to get cold, wet, or muddy,
depending on what we’re doing. Bring suitable footwear and clothing, and a pencil
and your lab notebook for data collection. For example, you must wear closed toed
shoes; for beach seining you must wear dive or surf booties, or old tennis shoes; for
some activities you might want to wear a wetsuit. If it rains or you get wet in the surf,
you must still be able to collect data. Field outings will last 3+ hours, so bring water,
snacks, and sunscreen.
Attendance:
This is an intellectually challenging and time-demanding course. You are expected to
attend and actively participate in every lecture, lab, and field experience. You will not
be able to make up missed lab or field activities.
III
Communication:
Outside of class time, we will communicate with you via your email account. If you
do not normally use this account, please configure your account to forward your
messages to your preferred account. You can expect us to respond to your email
within 24 hours (except on weekends – 48 hours). Your best source of course
information is the course website. Check the schedule on the website regularly
because occasionally we may have to change the schedule due to bad weather or other
unforeseen complications. Also, pdf files of required readings for lecture (scientific
papers), pdf’ s of PowerPoint lectures, class assignments, directions to field sites, and
data from our field studies are downloadable from the website.
Acknowledgements
We would like to thank Mrs. Noel for her contributions in sketching the original
scientific drawings, and the Academic Publication Council, Kuwait University for the
support in publication of the manual.
IV
TABLE OF CONTENTS
Preface & Acknowledgements I
Exercises
1. General characteristics, Taxonomy and Agnatha (Jawless fishes) 1
2. Gnathostomata (Jaw fishes) 27
3. Skin, Skin derivatives & Fins 42
4. Blood & its Circulation 57
5. Digestion 85
6. Age & Growth determination 104
7. Urinary system 119
8. Reproductive system 125
9. Fecundity measurement 138
10. Identification Key 147
11. References 157

Marine Ichthyology © Dr. Fadwa Abou Seedo & Dr. Subrahmanyam M N V
Exercise 1
1
GENERAL CHARACTERS
Fish are cold blooded vertebrates that breathe by means of gills and live in water.
They comprise about 30,000 to 40,000 species differing widely from each other from
shape, size, habits and habitats.
Some of them are very small, not more than an inch in length, while a few attain a
length up to 18.50 meters. They live in the seas, rivers, lakes, canals, dams and
almost all the places where there is water.
Fishes usually have a stream-lined body but some are elongated snake-like body and a
few are dorso-ventrally compressed body.
They have paired and unpaired fins supported by soft (or) spiny rays.
Dorsal, caudal and anal fins are unpaired while the pectorals and pelvic (ventrals) are
paired.
A number of species possess barbells which are excellent organs of touch.

They constitute economically a very important group of animals.
Besides being used as food, fish liver is an important source of oil contains vitamins A & D.
Body oil from fish is extensively used in soap industry and tanning.
Fishes also yield fish meal, fish manure, isinglass and several other products of
commerce.
It is, therefore, natural that man has paid a considerable attention to the study of the
anatomy, physiology and habits of the fishes. As compared to the tetrapods, fishes have
the following common characters, most of which are due to their permanently aquatic
life:
(i) Organs of respiration are gills.
(ii) Locomotory organs are the paired and unpaired fins which are supported by fin
rays of dermal origin.
(iii) An exoskeleton of scales is usually present but some species are naked
secondarily.
(iv) The notochord is partly replaced by cartilaginous or bony vertebrae.
2
(v) Paired visceral arches are present. The first pair forms the upper and lower jaws,
the second forms the suspensorium and the rest support the gills.
(vi) There is no middle ear.
(vii) A swim bladder is usually present (excepting in elasmobranches) but is
secondarily lost in some species.
(viii) Kidney is mesonephric.
3
TAXONOMY
 Fish are a paraphyletic group: that is, any clade containing all fish also
contains the tetrapods, which are not fish. For this reason, groups such as the
"Class Pisces" seen in older reference works are no longer used in formal
classifications.
 Traditional classification divide fish into three extant classes, and with extinct
forms sometimes classified within the tree, sometimes as their own classes
(Romer & Parsons, 1977)
 Class Agnatha (jawless fish)
o Subclass Cyclostomata (hagfish and lampreys)
o Subclass Ostracodermi (armoured jawless fish) †
 Class Chondrichthyes (cartilaginous fish)
o Subclass Elasmobranchii (sharks and rays)
o Subclass Holocephali (chimaeras and extinct relatives)
 Class Placodermi (armoured fish) †
 Class Acanthodii ("spiny sharks", sometimes classified under bony fishes)†
 Class Osteichthyes (bony fish)
o Subclass Actinopterygii (ray finned fishes)
o Subclass Sarcopterygii (fleshy finned fishes, ancestors of tetrapods)
 The above scheme is the one most commonly encountered in non-specialist
and general works. Many of the above groups are paraphyletic, in that they
have given rise to successive groups: Agnathans are ancestral to
Chondrichthyes, who again have given rise to Acanthodiians, the ancestors of
Osteichthyes. With the arrival of phylogenetic nomenclature, the fishes has
been split up into a more detailed scheme, with the following major groups:
 Class Myxini (hagfish)
 Class Pteraspidomorphi † (early jawless fish)
 Class Thelodonti †
 Class Anaspida †
4
 Class Petromyzontida or Hyperoartia

o Petromyzontidae (lampreys)
 Class Conodonta (conodonts) †
 Class Cephalaspidomorphi † (early jawless fish)
o (unranked) Galeaspida †
o (unranked) Pituriaspida †
o (unranked) Osteostraci †
 Infraphylum Gnathostomata (jawed vertebrates)
o Class Placodermi † (armoured fish)
o Class Chondrichthyes (cartilaginous fish)
o Class Acanthodii † (spiny sharks)
o Superclass Osteichthyes (bony fish)
 Class Actinopterygii (ray-finned fish)
 Subclass Chondrostei
o Order Acipenseriformes (sturgeons and
paddlefishes)
o Order Polypteriformes (reedfishes and bichirs).
 Subclass Neopterygii
o Infraclass Holostei (gars and bowfins)
o Infraclass Teleostei (many orders of common
fish)
 Class Sarcopterygii (lobe-finned fish)
 Subclass Actinistia (coelacanths)
 Subclass Dipnoi (lungfish)
 † Indicates extinct taxon
 The various fish groups account for more than half of vertebrate species.
There are almost 28,000 known extant species, of which almost 27,000 are
bony fish, with 970 sharks, rays, and chimeras and about 108 hagfish and
lampreys (Nelson, 2006).
 A third of these species fall within the nine largest families; from largest to
smallest, these families are Cyprinidae, Gobiidae, Cichlidae, Characidae,
Loricariidae, Balitoridae, Serranidae, Labridae, and Scorpaenidae. About 64
5
families are monotypic, containing only one species. The final total of extant
species may grow to exceed 32,500 (Nelson, 2006).
AGNATHA (Jawless Fish)
Agnatha are the earliest known vertebrates and are characterized by the absent of Jaws.
They are divided into several groups of which all excepting the "Cyclostomes” (living
Agnatha) are extinct and lived during the Silurian and Devonian times.
The fossil Agnatha are collectively known as "Ostracoderms" and were generally
heavily armoured (Fig. 1.1)
Living Agnatha are divided into the following classes:
Class: Myxini and Class: Cephalaspidomorphi
- The cyclostomes (Fig.1.2) are characterized by the presence of a suctorial mouth

(without functional Jaws), and a single nasal aperture and by the absence of paired
fins.
- They exhibit a low grade of organization.
They possess the following general characters:
(i) The body is elongated, eel-like and cylindrical.
(ii) The skin is smooth, slimy and without scales.
(iii) Respiratory organs are pouch-like (or) sac-like gills.
(iv) Mouth is circular and suctorial, without jaws.
(v) A few horny teeth and a rasping tongue is present.
(vi) Skeleton is cartilaginous, notochord persistent and a branchial basket is
present in the pharynyx.
(vii) There are no paired appendages. Generative ducts, sympathetic nervous
system and the conus arteriosus are not present.
(viii) Single nasal aperture and a median olfactory organ are present.
(ix) Alimentary canal is straight and there is no cloaca.
(x) Brain has a very small cerebellum. The membranous labyrinth has a single or
two semi-circular canals.
6
Class : Myxini
- These are commonly known as "hag-fishes" and are exclusively marine. The mouth
is terminal and the dorsal fin is absent (or) only feebly developed.
- The naso-hypophysial cavity opens behind into the pharynx.
- There are 1 to 15 gill apertures on each side, opening directly into the pharynx
- The branchial basket is reduced.
Examples: Myxine glutinosa, Bdellostoma (Eptatretus) cirrhatum, & Paramyxine etc.
Class: Cephalaspidomorphi
- This class includes the lampreys which possess one or two well developed dorsal fins
and a complete branchial basket.
- The naso-hypophysial cavity does not open behind into the pharynx.
- There are seven gill apertures on each side and they open into a respiratory tube lying
ventral to the esophagus.
Examples: Petromyzon marinus (sea lamprey), Lampetra fluviatilis (freshwater
lamprey), Ichthyomyzon & Mordacia etc.
GNATHOSTOMATA (Jaw Fishes)
"PLACODERMS"
- The 'placoderms' (Fig.1.3) were the earliest jawed vertebrates that appeared during
the "Silurian" and "Devonian" periods. Thus they lived during the 'Palaeozoic' era
only and are sometimes referred to as "Palaeozoic fishes".
- All of them were well protected by heavy, bony plates and are, therefore called
"Placoderms". All of them have few common features given below:
(i) All of them have a primitive type of Jaw. The upper jaw was strong and fused
with the skull.
7
(ii) The hyoid arch did not support the Jaws as in 'modern fishes', so that the Jaw
suspension was of a primitive type. This is called the "aphetohyoidean
condition".
(iii) Hyoid gill slits were present and 'spiracles' were not formed.
(iv) 'Paired' fins were present but were quite variable in different members of the
group.
Class : Chondrichthyes:
- This class includes a large number of marine fishes commonly called the sharks,
skates and rays.
- They were characterized by a cartilaginous exoskeleton which is partly calcified.
- The exoskeleton is in the form of dermal denticles or placoid scales.
- There are 5 to 7 gill slits on each side, opening directly to the exterior, operculum
being absent
- Tail is heterocercal and cloaca present.
- A spiracle is generally present.
- The skull is hyostylic and sometimes amphistylic.
- The males generally possess a pair of claspers on the ventral fins.
- Blood has a high concentration of urea.
- Air-bladder is not present and intestine has a spiral valve.
- Fertilization is internal and many species are viviparous.
- The elasmobranchii are mostly predatious fishes and are most abundant in tropical
& sub-tropical areas
- Some of them are very large in size and the biggest of all the fishes was probably
'Carcharodon' reaching about 80 feet in length.
- The Whale Shark, 'Rhinodon' is also the largest and reaches about 70 feet.
- The Basking shark 'Cetorhinus' grows to a length of 40 feet.
- Both these giant sharks are plankton feeders and are harmless to man.
However, a number of species like the Carcharodon carcharius (white shark)
Carcharinus glaucus (blue shark) and 'Tiger-shark' are man-eaters.
8
Class Chondrichthyes divided into 2 subclasses.

Subclass Elasmobranchii and Subclass Holocephalii
Sub-class: Elasmobranchii: (Figs. 1.4 &1.5)

Sub-class Elasmobranchii divided into 2 Superorders.
Superorder: Galea:
Ex: Horn sharks,
Collared carpet sharks,
Blind sharks,
Bamboo sharks,
Nurse sharks,
Zebra sharks,
Whale sharks,
Sand tigers,
Goblin sharks,
Crocodile sharks,
Mackeral sharks,
Megamouth sharks,
Thresher sharks,
Basking sharks,
Cat sharks,
Finback cat sharks,
False cat sharks,
Barbeled hound sharks,
Hound sharks,
Weasel sharks, and
Requiem sharks.
9
Superorder: Squalea:
Ex: Frill sharks,

Cow sharks,
Bramble sharks,
Sleeper sharks,
Dogfish sharks,
Angel sharks,
Saw sharks,
Guitarfishes,
Skates,
Electric rays,
Saw fishes,
Deepwater stingrays,
Longsnout stingrays,
River stingrays,
Stingrays,
Butterfly rays, and
Eagle rays.
Sub-class: Holocephali: (Fig. 1.6)
Ex: Plownose chimaeras,

Shortnose chimaeras, and
Longnose chimaeras.
Class : Osteichythes :
- This group includes a large number of 'bony fishes' in which the primary upper
and lower jaws are supplemented by the addition of membrane bones which form
the 'secondary jaws'. Hence, they are called the Teleostomi or perfect - mouthed fishes.
10
- The skull is usually hyostylie (or) 'amphistylic'.

- An operculum covering the gill-clefts is always present and there is a single
external branchial aperture.
- The body is generally covered with an exoskeleton of 'rhombic or cycloid scales.
- There is no cloaca, there being separate anal & urinogenital apertures.
- Claspers are not present
- Air-bladder is present in many species.
The Class Osteichthyes divided into 2 sub-classes, Actinopterygii and Crossopterygii.
I Sub-class: Actinopterygii (Ray fin fishes): (Today living fishes) (23,689 fishes approximately).
(Fig.1.7)
- This group includes the ray-finned fishes which are the commonest species of
today living in fresh water or sea.
- The fins of these fishes are not of the archipterygial type as their bony supports
are very much reduced and the fin-web is supported by fin-rays only.
- The internal nostrils are absent.
- The scales are well-developed and cycloid.
- Caudal fin is homocercal.
- Dorsal & anal fins are without spines.
- A spiracle is absent and an operculum covers the gill slits.
Sub-class: Actinopterygii is represented by Division Teleostomi.
Division: Teleostomi:
- This group includes a large number of bony fishes in which the primary upper and
lower jaws are supplemented by the addition of membrane bones which form the
secondary jaws.
- Hence they are called “Teleostomi” or ‘perfect mouthed fishes’.
- The body is generally covered with an exoskeleton of cycloid or rhombic scales.
11
- The gill filaments project beyond the inter-branchial septum which is greatly
reduced.
- An operculum covering the gill clefts is always present and there is a single
external branchial aperture.
II Sub-class: Sarcopterygii or Crossopterygii ( Lobe fin fishes): Eg: Latimeria sp

and Osteolepis
- Sarcopterygii (Fig.1.8) are characterized by the presence of paired fins having a
scale covered lobe.
- The pectoral fins are supported by an endoskeleton consisting of jointed median
axis bearing radials on each side.
- The scales are covered by a thick layer of cosmine.
- Internal nostrils were present.
- Branchiostegal rays are absent, replaced by jugular plates.
Sub-class: Crossopterygii is represented by Infraclass Dipnoi.
Infraclass : Dipnoi : (Fig.1.9)
- Dipnoi or the "Lung fishes" are the most interesting group of bony fishes and
show close affinity to the teleosts on the one hand, and to the "amphibians" on the
other.
- At present, Dipnoi are represented by three genera –
These are Neoceratodus (Australia)
Protopterus (Africa)
Lepidosiren (South America)
At present Dipnoi occurring in widely separated tropical & sub-tropical freshwater
habitats.
12
AGNATHA (JAWLESS FISHES)

Myxine (Hag-fish) and Petromyzon (Lamprey):
External Features:
- Petromyzon has an elongated eel-like, nearly cylindrical body with a laterally

flattened tail (Figs. 1.10 & 1.11).
- The mouth is situated on the ventral side of the anterior end, in the centre of a
funnel-like depression called the buccal-funnel.
- The margin of the buccal-funnel is beset with numerous papillae.
- A pair of large well-developed eyes is present, one on each side of the head.
- Seven pairs of small circular gill openings are present laterally behind the eye.
- A small, anal aperture lies on the ventral surface marking the junction between
the trunk and the tail.
- Paired fins are absent, but there are 2 median dorsal fins, the
caudal fin.
- The hag-fishes also have an elongated eel-like body but differ from lampreys in
many respects.
- The mouth is terminal and there is no buccal funnel.
- The mouth opening is surrounded by 4 pairs of tentacles supported by cartilage.
- Only one pair of external branchial apertures are present on the ventral side of the
body in Myxine, but in Bdellostoma there are 6-14 small gill openings on each
side of the body.
- The appendages are represented by a single caudal fin surrounding the tail and
there is no dorsal fin.
Skin:
- The skin of lamprey is slimy and is composed of several layers of epidermis.

Numerous unicellular glands are present in the epidermis and secrete large
amounts of mucous.
13
- In addition to these, hag-fishes have segmentally arranged 'thread-cells' forming

slime glands that open on the surface and secrete mucus.
Alimentary canal:
- In lampreys the buccal funnel is suctorial and armed with teeth.

- A pair of buccal glands opens below the tongue and secretes a substance which
prevents the coagulation of blood.
- The buccal cavity opens behind into two tubes, the dorsal one is the esophagus
and the ventral is the pharynx guarded by a velum. The esophagus is followed by
a straight intestine provided by a typhlosole, there being no stomach. The
typhlosole is called the spiral valve.
- Liver is a bilobed gland. The gall bladder and the bile duct disappear in the adult.
- There is no separate pancreas and the exocrine cells are embedded in the gut
epithelium.
- In Myxine there is no buccal funnel but the buccal cavity is longer than in
Lampreys.
- The intestine is wide.
- The liver consists of anterior and posterior lobes whose ducts open separately into
the gall bladder.
Respiratory organs:
- The respiratory organs consist of a number of branchial sacs with numerous

branchial lamellae on their inner surface.
- In Petromyzon, there are 7 pairs of these sacs and each opens to the exterior by an
aperture.
- Internally, the gill sacs opens into the pharynx which is in the form a sub-
esophageal tube, closed at the hind-end and is known as the respiratory-tube
- During respiration water enters into the branchial sacs through the external gill
apertures and is then expelled through the same route.
14
- In hag-fishes, six pairs of branchial sacs are present and open directly into the
esophagus by internal branchial tubes and by external branchial tubes.
- An oesophageo-cutaneous duct is present on the left side of the hag-fishes and
connects the oesophagus with the last gill sac.
- The respiratory water current, probably enters through the nasal tube and passes
out through the external branchial apertures.
Nervous system:
- The brain of Petromyzon is in a relatively primitive condition.

- The cerebellum is small, being represented by a mere transverse band.
- The optic lobes are imperfectly differentiated.
- The rest of the hind brain is well developed.
- The brain of Myxinoids on the other hand differs considerably from petromyzon
especially in the larger olfactory lobes, reduced ventricles and small midbrain.
- There are 10 pairs of cranial nerve in petromyzon, but in Myxinoids, the 3rd, 4th
and 6th appear to be absent due to the absence of eye muscles.
Urinogenital organs:
- The kidneys are very small in the hag-fishes.

- Pronephros persists, but the functional kidney is mesonephric.
- In petromyzon, the structure of the kidney is more complicated and pronephros is
not present in the adult.
- The sexes are separate in the adult lamprey but sexual differentiation occurs at a
late stage, so that both oocytes and spermatocytes may be present in the same
gonad in young lamprey.
- The gonad is unpaired. The ova as well as the sperms are liberated into coelom by
the rupture of the gonad.
- Sexes are separate in Myxine also, but the differentiation of the immature gonad
occurs late in life as in lamprey, so that a female may have an undeveloped testis
and vice versa.
15
PTERASPIS
HEMICYCLAPSIS
BIRKENIA
DREPANASPIS
Figure 1.1 Ostracoderms
16
PETROMYZON
MIXINE
BDELLOSTOMA
Figure 1.2 Cyclostomes
17
CLIMATIUS
COCCOSTEUS
BOTHRIOLEPIS
PTERICHTHYES
Figure 1.3 Placoderms (Extinct)
18
ELECTRIC RAY
EAGLE RAY
STING RAY
Figure 1.4 Elasmobranchii
19
SAW FISH
SKATE OR GUITAR FISH
BLUE SHARK
HAMMER HEADED SHARK
Figure 1.5 Elasmobranchii contd……
20
CHIMAERA
CALLORHYNCHUS
HARRIOTTA
Figure 1.6 Holocephali
21
ACIPENSER (STURGEON FISH)
AMIA (BOWFIN FISH)
ATRACTOSTEUS (GAR FISH)
Figure 1.7 Actinopterygii
22
LATIMERIA
OSTEOLEPIS
Figure 1.8 Sarcopterygii
23
DIPTERUS
NEO-CERATODUS
PROTOPTERUS
LEPIDOSIREN
Figure 1.9 Dipnoi
24
Figure 1.10
25
Figure 1.11 L.S. of Head region of Petromyzon
Figure 1.12 L.S. of Head region of Myxine
26
Exercise 2
27
GNATHOSTOMATA
Structure and Body Shape:
- The great ecological diversity of fishes is reflected in the astonishing variety of

body shapes and means of locomotion they possess.
- Indeed, much can be learned about the ecology of a fish simply by examining its
anatomical features or by watching it move through the water.
- Equally important to students of ichthyology is that these features also form the
basis of most schemes of classification and identification.
- Although the three main living group of fishes (jawless, cartilaginous, and bony)
share certain features, they differ considerably in several ways.
- The purpose of this exercise, therefore, is to provide an overview of external
anatomy.
- Using specimens available to you during laboratory sessions, locate the
morphological features described in the following paragraphs.
Cartilaginous fishes:
- The four groups of cartilaginous fishes (Chondrichthyes) have several distinguishing

features.
- Having a well developed lower jaw.
- The presence of true bony teeth on both jaws.
- In chimaeroids, the upper jaw is permanently fused to the cranium, while in the
shark, skates, and rays, it is separate.
- Other morphological features are paired fins, gills with individual openings, often
a pair of spiracles on the top of the head, and a pair of elongate claspers on the inner
edge of the ventral fins in mature males.
- Cartilaginous fishes have placoid scales, and these scales are developmentally and
histologically similar to teeth.
- Sharks have five to seven gill clefts on each side of the head and a free eyelid on the
upper portion of the eye. The body is divided into head, trunk, and tail regions, and
28
sharks have two sets of paired fins, distinct dorsal fins, often a median ventral fin,
and a heterocercal caudal fin with dorsal lobe larger than the ventral lobe.
- Skates and rays have greatly enlarged pectoral fins attached to the sides of the head
and the gills located in front of and below these fins.
- Skates and rays often take water in through the spiracles on top of the head instead
of through the mouth because they live on the bottom and their gills are oriented
ventrally. In addition, they lack a free upper eyelid.
Bony fishes:
- The bony fishes (Osteichthyes) are an incredibly diverse group, and it is impossible
to present all the morphological variations present in these fishes.
- There certain external features typical of bony fishes however, and we present
these as below:
Gross External Anatomy and Internal Support Systems of a Fish (Figs. 2.1 -2.7):
1. Body shape
2. Body covering (Skin)
3. Skin appendages
● Scales
4. Fins
● Paired fins
Pelvic
Pectoral
● Unpaired fins
Dorsal
Anal
Caudal
5. Openings
● Mouth
● Gill aperture
● Anus
● Abdominal pores
29
6. Sensory organs
● Nares (Nostrils)
● Eyes
● Skin organs
7. Skeleton
● Axial skeleton
Vertebral column
Skull
● Appendicular skeleton
Fins of internal support
Pectoral girdle
Pelvic girdle
8. Muscles
● Red muscles
● White muscles
● Pink muscles
1. Body shape:
- Commonly the fish body is torpedo-shaped (fusiform).

- In free-swimming species the body approximates the theoretically perfect
streamlined form.
- Many fishes depart moderately to completely from the foregoing generalized
shape.
- These departures range from globe shapes through serpentine, to thread-like
in outline.
- Some are strongly flattened from side to side, others flattened but greatly
elongated.
- And still others, flattened from top to bottom.
- In spite of the many variations in shape, the ground plan of body organization in
fishes is bilateral symmetry, as for the vertebrates generally.
30
2. Body covering (Skin):
- An ordinary fish is covered with a relatively tough skin.

- The skin is continuous with the lining of all the body openings and is transparent
as it runs over the surface of the eye.
- Much of the diverse coloration of fishes is due to its color cells, and the slimy
coating is due to its mucous cells.
- The skin in many fishes is devoid of scales, but in others scales that develop in it
armor it.
- Scales range in size from microscopic to large, in thickness from tissue-thin to
plate-thick, in ornamentation from simple to complex, in extend of body coverage
from partial to complete, in structure from non-bony to quite bony, and in fastness
from loosely deciduous to very firmly attached.
- The type of scales may characterize major fish groups.
- Scale morphology and the numbers of scale rows along or around the body
frequently serve specific and generic characters.
- Living agnaths are scaleless; sharks and their relatives have dentinal placoid
scales known as denticles; bony fishes have various types of bony scales.
3. Fins:
- Appendages of fishes comprise mainly, the fins. The fins are classified as
median or paired and unpaired.
Unpaired Fins:
- The rayed fins in line with the median axis of a typical fish are those of the back
(dorsal fin or fins), the tail (caudal fin), and the lower edge of the body just behind
the vent (anal fin).
- Although most commonly all of the foregoing are present, each is absent in some
kinds of fishes.
- Also developed in the median axis may be ray less, fatty adipose fin or fins
reduced to a few disconnected spines.
31
Paired Fins:
- Paired fins are the pectorals and the pelvic (ventrals).
- The pectorals are supported by the pectoral girdle that joins the skull and,
in spiny- rayed fishes, are situated higher on the sides of the fish than the pelvic.
- Pelvic fins vary substantially in position and in adaptive modification.
- Typically their support is by a pelvic girdle anchored in the belly musculature.
- In most soft-rayed fishes the pelvic fins are abdominal in position, but they may
also be situated anteriorly, just below the pectorals, in a thoracic position, or even
under the throat position.
4. Openings:
- The mouth, the gill apertures and the vent (anus) are the principal gross openings
connected with the alimentary canal in the fish body.
- Openings for the sense organs include the narial apertures and variously
distributed small sensory pores.
Mouth:
- The fish mouth is situated anteriorly on the head, in terminal position, but
adaptively its position may be superior (opening dorsally) or inferior (slightly to
prominently overhung by the snout).
Gill Apertures:
- In many fishes with gill covers there is a single opening on each side of the head.
- In the shark group, there is a spiracle that is located between the anterior most
gill slit and the eye. Inside the spiracle a small tuft of gill filaments (the hyoidean
P seudobranch) persists. This structure retained in the higher bony fishes
although they have no spiracle.
Anus:
- The anus or vent of a fish is on the mid-ventral line of the body. Most commonly
It is in the second half of the overall length of the individual, behind the bases of
the pelvic fins and just in front of the anal fin.
32
- In the sharks and their relatives, the anus opens into a depression on the ventral
body surface, the cloaca, which contains the exits of the urinary and genital
ducts, as well as that of the intestine.
Abdominal Pores:
- Abdominal pores open anteriorly in the vent of some fishes and sharks.
- These openings may not be identical in their origins in these two groups, but
in both they afford a communication between the body cavity and the exterior.
5. Sensory Organs:
Nares:
- One or two nares (nostrils) on each side of the snout leading to a blind sac
represent the organs of smell externally among fishes.
- Most fishes have the narial openings at the top and sides of the snout.
- In others, such as sharks, rays and skates the nares are on the ventral
surface of the snout.
Eyes:
- The essentially lidless eyes that cannot be closed are situated in orbits, one
on each side of the middle line of the fish head.
- Most often the eyes are lateral, with partially independent fields of vision
and movement.
Skin:
- Numerous microscopically small openings of skin sensory organs are developed
on the surface of the fish body.
- In most fishes a series of these pores, extending along each side in a single row
from the head to the caudal fin, comprises the lateral line.
- The lateral-line system forms branches about the head, including one above and
one below the orbit of each eye.
33
6. Skeleton:
Upper Jaw:
- The Jaws have undergone rather dramatic changes in bony fishes, related to the
change from a firm biting mouth to a flexible sucking mouth.
- In the course of this change, the principal bones of the upper jaw have changed
from the maxillas to the premaxillas.
- The principal bones bearing teeth have also changed.
Lower Jaw:
- In contrast with the upper jaw, the principal bones of the lower jaw (dentaries)
have remained fairly constant.
Opercular Bones:
- The opercular bones have also remained fairly constant in bony fish, although
some have added an interopercular bone on each side to increase the efficiency
of the operculum as a respiratory pump.
7. Muscles:
- In almost all fish, the large muscles of body and tail comprise the majority of the
body mass, although there are many other muscles associated with the head and
fins.
- The body muscles are divided vertically along the body length into sections, the
myomers (myotomes), which are separated by sheets of connective tissue.
- The myomers are shaped like a "W" on its side, so that they fit into one another
like a series of cones. The myomers on the right and left halves of the body are
separated by a vertical septum.
- A horizontal septum separates the muscle masses on the upper and lower halves
of the body. The upper muscles are called the epaxial muscles and the lower
muscles the hypaxial muscles.
- In addition, there is usually a lateral band of muscles that lies along or slightly
below the midline of the fish. On inspection, fish muscles can often be divided
into red ( slow), white (fast), and pink (intermediate) muscle.
34
Figure 2.1 Some meristic and morphometric characters

shown on a hypothetical marine fish
35
Figure 2.2 Variations in body form among fishes
36
Figure 2.3 Typical fish body shapes

(A) and (B) rower-predator; (C) lie-in-wait predator;
(D) surface-oriented fish; (E) bottom rover;
(F) bottom clinger; (G) flatfish; (H) rattail
(I) deep-bodied fish; (J) eel-like fish
37
Figure 2.4 Comparative external anatomy of major

groups of living fishes
38
Figure 2.5 External features of a bony fish (snapper, top) and a

cartilaginous fish (smooth hound shark, bottom)
39
Figure 2.6 A Diagram of a siluroid fish (top);

B Diagram of a scaly fish (bottom)
40
Figure 2.7 External features of a shark (side view, above);

and a skate (top view, below)
41
Exercise 3
SKIN
- The skin forms the external covering of the body and performs a number of
important functions in fishes.
- The skin contains sensory receptors tuned to the surroundings of a fish.
- Besides protecting the body against injury and infections, the skin has respiratory,
excretory & osmoregulatory functions.
- The derivatives of the skin play an important part in the metabolic activities of the
body. Special structures of some species like the electric organs, poisonous
glands, and phosphorescent organs are the integumentary derivatives.
Structure: (Figs. 3.1 A&B)
- The skin of a fish is composed of 2 layers, an outer epidermis & an inner

dermis or corium.
- The epidermis is ectodermal in origin and consists of several layers of flattened
cells (stratified squamous epithelium), of which the deepest layers are made up of
columnar cells forming the stratum germinativum in which cells are always
multiplying by mitotic division to replace the outer worn out cells.
- A superficial layer of dead horny cells, forming the stratum corneum in terrestrial
vertebrates, is not present in fishes.
- Numerous tubular or flask shaped mucous cells are scattered among the epidermal
cells and may even extend into the dermis.
- The cells secrete mucin, a glycoprotein, which mixes with water to form a thick,
slimy mucous, covering the whole body.
- Mucous cells develop from the stratum germinativum and migrate to the surface.
They vary in number, kind and size in different species of fishes.
- The dermis is mesodermal in origin and is composed of connective tissue, blood
vessels, nerves and cutaneous sense organs.
- The thin upper layer of loose connective tissue is called the stratum spongiosum
and the thicker dense lower layer, the stratum compactum.
43
Functions:
- The mucus covering the body of the fish, performs several important functions.
- It lubricates the fish so as to reduce body friction in water while swimming. Thus
enabling the fish to move with a greater speed.
- It protects the body from parasites, fungus, bacteria and other microorganisms.
- In turbid or muddy water, the mucus has the power to precipitate the particles in
suspension especially in contact with the gills and thus prevents the fish from
suffocation and helps in its survival.
- The skin & the mucus secreted by its glands help the fish in regulating to some
extent, the osmotic changes of water and ions between the body fluids and the
surrounding medium.
- The skin performs an important function in repairing the surface wounds.
Immediately after an injury, the wound is closed by the mucus containing a large
number of lymphocytes.
- The skin is also an important respiratory organ, the fish (certain species) are able
to survive for a considerable time out of water due to highly vascular dermis of
their skin.
- Glandular cells of the epidermis are modified to form poison glands in certain
fishes and are used for offence as well as defense.
- Light producing phosphorescent organs of a number of marine fishes are also the
derivatives of the epidermis.
- Chromatophore of various kinds, present in the dermis, gives various colour
patterns to the body, making it conspicuous or inconspicuous.
- Skin has also the power to absorb dissolved nutrients from the surrounding water.
44
SKIN DERIVATIVES
SCALES (Figs. 3.2, 3.3 & 3.4)
- In most of the fishes, the skin is covered with an exoskeleton in the form of scales
and only a few are naked having no scales on the body as in cat-fishes.
- According to the mode of their origin, there are 2 types of scales:
(i) those which are formed due to the secretory activity of both epidermis &
dermis as the PLACOID scales of elasmobranches, and
(ii) NON-PLACOID scales that are derived from the dermis only as the scale
of teleosts. Structurally, the scales are classified as COSMOID, GANOID
(or) RHOMBOID, PLACOID, CYCLOID & CTENOID, the last two are
also called as the bony ridge scales.
PLACOID
- Placoid scales are dermal denticles and is characteristic of the sharks.

- Each has a disc-like basal plate embedded in the dermis and a spine projecting out
through the epidermis. In structure, placoid scale resembles a tooth.
- The spine has an external covering of enamel-like, hard, transparent material
called vitrodentine. This is followed by a layer of dentine enclosing a pulp cavity
from which several branching dentinal tubules radiate in different directions.
- The centre of the basal plate is perforated by an aperture to provide entrance to
the blood-vessels and nerves from the dermis.
- The scales are closely set but do not overlap each other.
COSMOID
- The cosmoid scales are found in the extinct Crossopterygii and Dipnoi (lung
fishes). The external layer of the scale is thin and enamel-like and is called the
vitrodentine.
- The middle layer is made up of hard, non-cellular, dentine like material called the
cosmine, and contains a large number of branching tubules and chambers.
45
- The inner layer is made up of vascularised bony-substance, isopedine.

- These scales grew by the addition of new isopedine material from below, along
the edges.
- Cosmoid scales are not found in the living fishes.
GANOID (or) RHOMBOID
- These are characteristic of the primitive Actinopterygians, and are various forms
and structure.
- These scales are heavy and have an outer layer of hard inorganic, enamel-like
material called ganoine.
- The middle layer is cosmine containing numerous branching tubules.
- The innermost layer is thickest and is made up of lamellar bone, isopedine.
- These scales grow by the addition of new layers to lower as well as upper surface.
- These scales are usually rhomboid in shape and articulate by peg & socket joints.
BONY RIDGE (Cycloid & Ctenoid)
- They are present in most of the teleostean fishes, and are thin, flexible, transparent
structures due to the absence of the first and middle layers of other types.
- These scales exhibit characteristic ridges alternating with grooves and generally
the ridges are in the form of concentric rings. The central part of the scale is
called the focus and is the first part to develop.
- In many species, oblique grooves (or) radius run down the focus towards the
margin of the scale.
- The origin & development of both these types of scales are thin and roughly
rounded in shape, being thicker in the center and thinning out towards the margin.
- The cycloid scales are round, flat & thin.
- In cycloid scales, the posterior field may often be identified by ridges, are least
distinct there, often appearing as if they had been eroded or abraded.
- Pigment cells (chromatophores) adhere in this zone.
46
- The ctenoid scales are found mainly on spiny-finned teleosts.

- The ctenoid scales are characterized by the tiny, comb-like projections (ctenii or
teeth) on the exposed (posterior) edge of the scales.
47
FINS
- Fins are the chief organs of locomotion in fishes, and are of 2 kinds, Paired
fins and Unpaired fins.
- The paired fins are the pectorals and pelvics corresponding to the fore & hind
limbs of terrestrial vertebrates.
- The Unpaired fins include a dorsal on the back, an anal on the ventral side
behind the vent and a caudal at the end of the caudal peduncle.
- Both the paired and unpaired fins are supported by skeletal rods called the
radials and dermal fin rays.
- In teleosts the fin rays are branched and jointed by structures are called the
lepidotrichia.
- In sharks & rays the fin rays are called ceratotrichia and are fairly stiff,
unbranched and unsegmented.
PAIRED FINS (Fig. 3.5)
- Paired appendages were not present in the ancestral vertebrates and were
developed during the course of early fish evolution.
- The supporting endo-skeleton of the paired fins varies greatly in different
groups of fishes.
- The original primitive condition resembled the fins of Ceratodus and is known
as the 'archipterygium'. It had a median jointed axis articulating with the
girdle and was provided with an anterior pre-axial and a posterior, post-axial
series of radials attached to it. The radials were arranged on either side of the
median axis and decreased in size towards the tip. This is called the biserial
arrangement of radials. This type of "biserial archipterygium" is found in
Ceratodus only.
- Various types of paired fin skeleton in the Teleostomi may be derived from the
archipterygium by shortening of the median axis and reduction in the number
of post-axial radials that finally disappeared completely, giving rise to
'pleuroarchic' (or) uniserial type of skeleton found in teleosts.
48
UNPAIRED FINS (Figs. 3.6 & 3.7)
- The unpaired fins of all the fishes develop as a result of differentiation in a

continuous embryonic fin-fold.
- In higher fishes, separate dorsal, caudal & anal fins are formed by the
concentration of radials in certain areas and degeneration of the fold in the
intervening spaces, between the fins.
- The dorsal and anal fins are provided with a fleshy lobe at the base consisting
of fin muscles surrounding the basals and the radials.
- In all the higher bony fishes, the fleshy lobe at the base of the fin has
disappeared and the radials are reduced to nodules of bone (or) cartilage.
- Caudal fin differs from the dorsal & anal fins in the nature of its supporting
skeleton and is of 3 main types:
(i) The Protocercal (first tail) is probably the most primitive. Here, the hind end
of the notochord or the vertebral column is straight and divides the caudal fin
into two equal lobes, the dorsal epichordal, and the ventral hypochordal lobe.
The term 'diphycercal' is also used for secondarily symmetrical tails as in
Dipnoi but the term 'protocercal' is preferred for all types of symmetrical tails.
Eg: Amphioxus & Cyclostomata.
- In all the fishes, the caudal fin passes through the protocercal stage during
development.
(ii) The Heterocercal (unequal) tail is characteristic of the chondrichthyes and
some primitive bony fishes.
- Here, the hind end of the notochord is bent upwards and continues almost upto
the tip of the caudal fin.
- The ventral hypochordal lobe is much larger than the dorsal epichordal, so that
the caudal fin is asymmetrical both externally and internally.
(iii) The Homocercal (equal) tail is characteristic of most higher bony fishes.
- It is symmetrical externally consisting of equal sized epi-, and hypochordal
lobes. But internally the tail is asymmetrical and the hinder end of the
vertebral column is turned up-ward and greatly shortened. The end of the
vertebral column does not reach the posterior limit of the fin.
49
- Actually, the epichordal lobe is much smaller than in the heterocercal tail, as
almost the entire caudal fin of the homocercal type is derived from the lower
lobe, and the upper lobe contributes only a few, small, unbranched rays to the
upper margin of the fin.
- The homocercal tail is derived from the heterocercal type and intermediate
types are found in many bony fishes. The transition is seen in many fossil
bony fishes.
50
Figure 3.1
A. Structure of a fish skin without scales
B. Structure of a fish skin with scales
51
Figure 3.2 Kinds of scale shapes
52
Figure 3.3 Location of the first scales on the body of different types of
fishes
53
Figure 3.4 Examples of different kinds of scales on different fish types
54
Figure 3.5 Origin of paired fins
55
Figure 3.6 Classification of caudal fin on the basis of supporting

internal skeleton
56
Exercise 4
BLOOD AND ITS CIRCULATION
All fish tissues require a delivery of nutrients (including oxygen) to function properly.
The circulatory system, with blood as its medium, delivers these essential nutrients
throughout the body and removes both metabolic wastes and pathogens before
harmful concentrations are reached.
BLOOD:
Fish blood, like that of other vertebrates and many invertebrates, is composed of
blood cells suspended in plasma that are circulated throughout the body tissues.
Blood Volumes:
- Range from 3% to 7% of body weight in bony fishes
- Range from 4% to 8% of body weight in elasmobranchs.
- Range from 8% to 2% of body weight in agnathans.
Blood cells are of two basic types:
- Erythrocytes ( red blood cells or RBC’)
- Leucocytes ( white blood cells or WBC’s)
Erythrocytes ( RBC’s):
- Mature erythrocytes are oval, tiny & nucleated.
- RBC’s are the most abundant cells in fish blood ( up to 4 million cells/ mm3 )
- RBC’s have been found in the blood of all fishes, except in three small
Antarctic species a low, cold adjusted metabolism and inhabit oxygen rich
waters, and in the ribbon-like Leptocephalus larvae of eels ( Anguilla ) and
certain deepsea fishes.
- RBC’s contain haemoglobin and carry oxygen from the gills to the tissues.
- Elasmobranchs typically have RBC’s that are larger, although fewer in
number, than those of teleosts.
58
- Teleosts, speices having more erythrocytes per milliliter of blood generally

have smaller RBC’s within this group, the more active species tend to have
more RBC’s than do sedentary species.
Leucocytes (WBC’s):
-
WBC’s are less abundant (20,000- 150,000 cells/ mm3) than RBC’s in fish
blood.
-
WBC’s are ovoid to spheroid in shape.
-
The principal types of WBC’s are
a – Lymphocytes
- Vary in size (diameter, 4.5-12 µm)
- They are dominated by the nucleus, with only a narrow rim of
basophilic cytoplasm.
- Their primary function, antibody production.
b- Thrombocytes
- Function in the clotting of circulatory fluid.
c- Monocytes/ macrophages
- Comprise a small proportion of the WBC.
- Function in phagocytize foreign particles.
- Morphologically, the cell outline may be quite irregular.
d- Granulocytes
- Having conspicuous cytoplasmic granules. Three basic types are found.
(1) Eosinophils, (2) Basophils and (3) Neutrophils
- The names of the three types reflect the comparative visibility of the
granules when the cells are treated with acidic, basic and neutral stains
respectively.
- Neutrophils are the most abundant, and often have eccentric nucleus,
which divided into several lobes.
- Granulocytes are formed in the kidney and spleen in teleosts, and in the
leydig organ in elasmobranches.
59
CIRCULATION:
The primary circulatory system of most fish is a closed system that typically
consists of single heart as the pump in line with branchial (gill) and systemic
(body) capillary beds connected by arteries and veins.
Heart structure:
- Blood circulation of most fishes is accomplished by a four-chambered heart in
line and pump venous blood. Except for air-breathing fishes, all flow of blood
is to the gills
- The heart and gills are closely associated, because fish hearts are located the
farthest anterior relative to those all the vertebrates.
- The heart is enclosed in the fluid –filled pericardial cavity, which is separated
from the peritoneal cavity by the pericardial wall, which is more rigid in
elasmobranches than in teleosts.
- The first chamber of the fish heart is sinus venosus, thin –walled chamber,
receives vinous blood from liver and ducts of cuvier.
- Venous (oxygen poor) blood directed to the second chamber atrium, through
sinoatrial valve.
- The atrium is a relatively large chamber that empties ventrally to the ventricle,
via atrioventricular valve (third chamber)
- The ventricle also is a relatively large chamber featuring thick walls of cardiac
muscle. It is pyramid- shaped in elasmobranhs and conical in teleosts, with
apex pointing posteriorly.
- Ventricular walls may be composed of two layers of muscle, the cortex and the
spongy myocardium.
a) The cortex is relatively a dense cardiac muscle (compact myocardium) that
generally receives oxygen and nutrients from the coronary artery.
b) The spongy myocardium, consists of a spongy mesh that is supplied with
oxygen and nutrients only by the venous blood that it pumps.
- The fourth chamber conous arteriosus in elasmobranchs, lamprey, hagfish,
and holosteans, and bulbus arteriosus in teleosts. The conous have many
valves aswell as cardiac musculature in the walls.
60
Blood vessels :
- The blood vessels in the blood of a fish are principally of two kinds:
- (a) the major arteries and veins, which follow the long axis of the body,
(b) branches of the major vessels, which lead to and from the skin, the
skeleton, musculature, and which also provide circulation for the brain, the
spinal cord, and the intestinal organs.
- Blood from the tail collects principally in the caudal vein as it passes forward
through the haemal arches just beneath the caudal artery.
- The caudal vein drains into the renal portal system of the kidneys and upon
leaving the kidneys joins blood from dorsal musculature in posterior cardinal
vein.
- Cardinal veins join the duct of cuvier (common cardinal veins) on each side of
the esophagus, near the level of the pectoral girdle.
- The ducts of cuvier receive venous blood from head and from the lateral body
Wall.
- A branch of the afferent artery leads into each gill filament where it breaks up
into capillaries in the gil lamellae of the filament.
- The oxygen rich blood from the filament collects into an efferent loop around
each branchial aperture.
- Efferent gill arteries merge to become dorsal aorta.
- In the lampreys and elasmobranch group of fishes, the efferent branchial
vessels rise dorsally one by one to join the median aorta.
- In higher bony fishes, the efferent branchial arteries of each side rise to collect
into a trunk vessel on each side. The result is that, over the gill region the
dorsal aorta is paired and does not become a median vessel until just behind
the gill region.
- The ventral aorta runs forwards and gives off four pairs of afferent branchial
vessels of which most bony fishes shows common root for 3rd and 4th gill
arteries.
- A special arrangement is found in the blood supply to the spiracle of sharks,
rays and skates and to its homologous structure in the bony fishes, the
hyoidean pseudobranch.
61
- Both are supplied with oxygenated blood from mandibular artery or its
derivatives.
- In bony fishes, an efferent pseudobranchial leads to the blood that has passed
the pseudobranchial capillaries through the ophthalmic artery to the choroid
gland of the eye.
GILLS:
- One of the fundamental needs of a fish, like other animals, is to have an

adequate supply of oxygen in the tissues, so oxidation can occur and provide
the necessary energy for life. The success of fish depends upon its ability to
obtain oxygen from the external environment by means of vascularized gills,
lungs or skin; to transport this oxygen to the tissues; and to unload the oxygen
to the tissues.
Structure of gills:
Lampreys:
- All lampreys have seven pairs of gill sacs or branchial pouches.
- The gill sacs open towards the lumen of the alimentary tract.
- Each gill sac divided from the next by thin diaphragm that adheres to the body
wall.
- The inside of each gill sac is covered with radially arranged gill filaments
which have small secondary cross folds on them to enlarge the respiratory
surface except the inner edge of the diaphragm.
- The branchial pouches supported by cartilaginous branchial basket of lattice-
work structure communicate with the exterior through more or less
pronounced, epithelium-lined branchial atria that open to exterior through the
gill pores or external branchial apertures.
- Between each branchial atrium and its external gill pore there is a short,
oblique, posteriorly directed branchial canal.
- Lampreys rarely use the suctorial mouth for inspiration, even when the mouth
is not attached to an object or a host fish.
- Fifty to seventy contractions of the gill pouches have been counted per minute
in the sea lampreys attached to prey, but have been found to rise to hundred
62
and twenty or even to two hundred contractions per minute in the rapidly
swimming river lampreys.
Hagfish:
- Hagfishes gill structure like lampreys are in pouches. Each pouch has an
internal opening from the pharynx and one that leads to or toward the outside.
- Along the sides of the body are from 1 to 14 gill openings.
- The hagfishes have two distinct breathing habits.
a) When the animal is not feeding, it may lie buried in the mud, except for the
anterior part of the head. When the animal in this position, water reaches the
gill pouches from the nasopharyngeal cavity.
b) When the mouth is buried in food, respiratory water flows alternately in and
out of the gill pouches through the esophageo-cutaneous duct. This duct opens
externally behind the last gill pouch.
Sharks & Rays:
- Five, more rarely six or seven , external gill slits exist ventrally in series on
each side in rays and their relatives, but laterally in sharks.
- An anterior to this series of openings in both the sharks and rays is the
spiracle, corresponding to vestigial primitive first gill slit.
- Well developed septa with cartilaginous supports and individual gill-arch
muscles characterize each holobranch, the oral and aboral sides of the septa
each carry a hemibranch composed of the gill tissue proper.
- Both primary and secondary gill filaments are detached from the septum, so
that two hemibranchs in opposition may form an effective barrier that forces
the water to penetrate between all filaments when it seeks it exit as a result of
the suction and pressure of respiratory movements.
Bony fishes:
- Bony fishes in general share the same basic arrangement of a single external
branchial aperture on each side of the head.
- Bony fishes don’t have spiracle, and they have four gill slits.
- Characterize by deepening indentation between the two hemibranchs of each
holobranch form a transition from lower (chondrichthyes) to higher (teleosts)
rayfin bony fishes.
63
- The septum between two opposed filaments may have become shortened
almost to the point of disappearance.
- When septum is reduced, opposed filaments are free from one another
throughout all or most of their length, depending on the extent of septal
reduction.
- The forked efferent branchial artery of each aortic arch of sharks is replaced
by single efferent vessel in the teleosts.
- In all bony fishes, muscles move the bases of the hemibranchs so that the
pressure in, and possibly also the flow of water from, the oral cavity is
regulated.
-
PSEUDOBRANCH:
- Although lacking a spiracle, ray fin bony fishes retain a hyodean pseudobranch
that is free in some fishes but skin- covered in others.
- It develops quite early in the embryo and may have a respiratory function
in the embryonic stage but not in adult.
- It received oxygenated blood directly from the dorsal aorta and has a vascular
connection with the internal carotid artery.
- It may serve to increase the oxygen concentration in the blood going to the
brain and the eye through the internal carotid artery.
- Each filament of a pseudobranch has a thin cartilaginous supporting rod and
blood vessels in its core, around which are arranged layers of pseudobranch
cells and sinusoidal blood spaces.
- Histologically, the pseudobranch contains a large number of acidophilic
secretory cells which probably produce carbonic anhydrate- like enzyme.
- The pseudobranch may also be useful in the filling of gas bladder and in the
regulation of intracellular pressure.
64
SWIM BLADDER:
- The swim bladder less precisely designed as the “gas bladder” or the “air
bladder” is characteristic of bony fishes and reaches its fullest development as
hydrostatic organ among the spiny-rayed teleosts.
- Swim bladder are of two basic types: physostomous and physoclistus.
- Physostomous swim bladders have connection (pneumatic duct) between the
swim bladder and the gut but in physoclistus swim bladders lack this
conection.
- Fish with physostomous swim bladder include many of the more ancestral,
soft-rayed teleosts, including herring, salmonids, osteoglossids, mormyrids,
pikes, cyprinids, characins, catfishes, and eels.
- Physostomous fish inflate their swim bladder by gulping air at the water
surface and then forcing it through the pneumatic duct and into the swim
bladder by a bucal force mechanism.
- In contrast to physostomous fish, fish with “closed” (physoclistus) swim
bladders have special structures associated with the circulatory system for
inflating or deflating the swim bladder.
- More than two-thirds of all teleosts (especially spiny-rayed species) are
physoclistus.
- Arete mirable (‘wonderful net”) and associated gland are the source of
inflation gas in these swim bladders
- The swim bladders functions also as an (a) accessory breathing organ,
(b) sound reception organ, (c) resonator in sound production organ.
65
Figure 4.1 Comparative generalized schemes of the heart in

major groups of living fishes
66
Figure 4.2 Arrangement of blood vessels in the respiratory region
67
Figure 4.3 Afferent and efferent branchial arteries of shark
68
Figure 4.4 A. Heart of bony fish

B. Heart to show internal structure
C. T.S. of auriculo-ventricular aperture
showing two pairs of valves
69
Figure 4.5 Diagrammatic cross section of teleost (Osteichthyes) and

shark (Chondrichthyes) showing the heart chambers,
valves and root of blood flow
70
Figure 4.6 Arrangement of major veins
71
Figure 4.7 The structure of fish blood vessels

A. Part of the wall of an intestinal artery
B. Arteriole
C. Wall of an intestinal vein
D. Capillary
Ad. adventia, CF. collagen fibres, DCF. dense collagen fiber, EF. Elastic fibres
End C. endothelial cell, IEL. Internal elastic lamina, In. intima, Md. Media
SMC. Smooth muscle cell
72
Figure 4.8 Blood cells of fishes

(a) erythrocyte, (b) lymphocyte
(c) monocyte, (d) thrombocyte
(e) acidohilic granulocyte,
(f) neutrophilic granulocyte
(g) basophilic granulocyte
73
Figure 4.9
(a) Gill arches of fishes
(b) T.S. Gill of a bony fish
74
Figure 4.10 Structure of gill arches
75
Figure 4.11 Types of gill structure
Figure 4.12 Ventilation in teleost fish
76
Figure 4.13 Generalized diagram of gill circulation (left side)

A. Shark group (Chondrichthyes)
B. Bony fish (Osteichthyes)
77
Figure 4.14 Horizontal sections through gills of

A. Shark
B. Bony fsh with detailed cross section
C. Near apex of B
78
Figure 4.15 Physostomous swim bladder (Open swim bladder)
Figure 4.16 Physoclistus swim bladder ( Closed swim bladder)
79
Figure 4.17 Different modifications of swim bladder in Osteichthyes
80
Figure 4.18 Types of swim bladder in teleost fishes
81
Figure 4.19 Swim bladder with T-shaped septum and alveolus
82
Figure 4.20 Extension of swim bladder to regions of inner ear and anus
83
Figure 4.21 Position of the pseudobranch
84
Exercise 5
DIGESTIVE SYSTEM
Feeding habits according to diet:
Fish can be classified broadly on the basis of their feeding habits as detritivores,
herbivores, carnivores, and omnivores. Within these categories fish can be
characterized further as (1) euryphagous, having a mixed diet; (2) stenophagous,
eating a limited assortment of food types; and (3) monophagous, consuming only one
sort of food. A majority of fish, however, are euryphagous carnivores.
.Major feeding types:
- As for the manner of feeding, only one broad common characteristic

prevails – the food is taken into mouth.
- Other than this the feeding habits and adaptations of fishes are very diverse.
- Nevertheless, certain very broad types of food getting are recognizable either
by species or by life history stages.
- On this basis, fishes can be classified, although somewhat arbitrarily, accord-
ing to their feeding behaviour as predators, grazers, food strainers, food
suckers, and parasites.
Feeding Adaptations:
The diversity of feeding habits that fishes exhibit is the result of evolution leading
to structural adaptations for getting food from the equally great diversity of
situations that have evolved in the environment.
86
i.) Modifications in the Lips: (Fig. 5.1.A)
- A significant advance in vertebrate evolution was the appearance of true jaws

to border the mouth opening.
- Most generally the jaw-equipped mouth has a biting function and fishes that
swallow large morsels of food usually have unmodified relatively thin lips.
- Suctorial feeders have an inferior mouth and fleshy modification of lips.
- Many suctorial feeders also have well developed barbels more or less
bordering the mouth.
- The barbells have many sensory end organs and help to locate food grubbed
from soft bottom materials.
- The suctorial lips of free-living fishes also serve as holdfast organs in fast
flowing mountain streams.
- Some fishes with suctorial lips adaptation, the opercular structure with a
separate water inhalant and exhalent device.
- Fishes with small mouth and inner surface of the lips has rasp-like folds
facilitate scraping of algae from the stones to which the fish adheres.
- In parasitic members of lamprey family, the jawless suctorial mouth serves
both as hold-fast for attachment to the host and as a food remover from the
host. Sucking disc of lamprey is also used to dislodge and transport stones
from the nest pit in stream.
ii.) Modifications in the shape of the mouth: (Figs. 5.1.B & 5.2)
- Among the grazers and suctorial feeders there exist not only specially develop-
ed lips but also adaptations of other mouth parts.
- Some fishes have mouths that resemble elongated beaks. This adaptation is
achieved by a protraction of the hyomandibular bone rather than by a
lengthening of the lower jaw bones (dentaries) themselves.
- Among the fishes, the method of feeding may be suction such as a
syringe, or it may be selective grazing action with sharp teeth where the long
snout enables to reach into small crevices of the coral reefs.
87
- Some predator fishes can form temporary tubes in which to engulf their prey
from close range by forward extension of the jaws enabled by special
- articulation of the premaxillaries and other skull bones.
- A peculiar structure among mouth modifications has arisen in the halfbeaks
where the lower jaw projects into a beak, often a third of the length of the fish
itself, with the mouth opening above it.
iii.) Modifications in the Teeth: (Fig. 5.3)
- Outstanding among the obvious adaptations for feeding in fishes are the teeth.
- They are thought to have arisen from scales covering the lips, as represented in
sharks, where the placoid scales of the skin visibly grade into teeth on the
jaws.
- In the bony fishes, teeth are of three kinds, based on where they are found:
jaw, mouth, and pharyngeal.
- Jaw teeth are variously those on the maxillary and premaxillary bones above,
and on the dentaries below.
- In the roof of the oral cavity, teeth are variously borne by the median vomer
and by the palatine and ectopterygoid bones on each side.
- In the floor of the mouth, the tongue often has teeth on it.
- Pharyngeal teeth occur as pads on various gill arch elements in many
species.
- Based on their form, some major kinds of jaw-teeth are the following:
cardiform, villiform, canine, incisor, and molariform.
- Cardiform teeth are numerous, short fine , and pointed.
- Villiform teeth are more or less elongated cardiform ones, in which the
length- to- diameter relationship resembles that of intestinal villi.
- Canines are dogtooth –like, often even quite fang-like, which are elongated and
subconical straight or curved.
- Incisors are sharply edged cutting teeth, some can be crenulated and in other
cases become fused into cutting beaks.
- Molariform teeth are for crushing and grinding, and hence have flattened with
88
broad surfaces.
iv.) Modifications in the Pharyngeal teeth: (Figs. 5.4 & 5.5)
- These consists of two sets of bones, a complex of three in the roof of the
pharynx and a pair on its floor.
- The members of the lower pair always united firmly into single bone, usually
more or less triangular in shape, whose apex is directed forward.
- The position of the upper and lower pharyngeal bones at the back of the throat
is such that all the food must pass between them.
- Both upper and lower pharyngeal bones are movable, the upper bones slide
mostly forward and backward, and to lesser extent side to side, whereas, the
movement of the lower bones is upward and downward, that is, towards and
away from the upper bones.
- Combination of these movements allows a wide variety of “processing”
operation to be performed which are capable of crushing and tearing food
and of raking into the esophagus.
- The pharyngeal pad (or palatal organ), which situated dorsally at the entrance
of the esophagus has been implicated in removing excess water from the
ingested food ( Ex. common carp).
- Pharyngeal valve, hangs from the roof of the pharynx of parrot fish, probably
assists in the placement of pieces of coral for grinding by pharyngeal teeth and
lubrication from epithelial mucous cells.
v.) Modifications in the Gill rackers: (Fig. 5.6)

- Gill rackers are forward-directed projections from the inner margin of gill
arches, guarding the internal branchial openings.
- Besides protecting the tender gill filaments from abrasion by ingested materials
that are coarse in texture, gill rackers are also specialized in relation to food
and feeding habits.
- They are very stubby and unadorned in omnivore fishes.
- In many plankton feeders, the gill rackers are elongated, numerous, and
and variously lamellated or ornamented, presumably to augment efficiency in
89
straining.
- Simple, but very numerous rackers and ornamented structures are however
found in certain fishes and are of taxonomic use.
- In piscivorous fish (Ex. large mouth bass), the gill rackers are short, stout,
widely spaced and pointed. Preventing the prey from escaping through (and
damaging) the gills, but they also may partially descale the prey as it passes
by the rackers.
- Shortest gill rackers are specialized for fish feeding on snails.
- Gill rackers of intermediate length, thickness and spacing are found in
euryphagous species.
- The largest and finest gill rackers, reflect species of zooplankton in their diets.
(vi) Modifications in the Digestive tract: (Figs. 5.7, 5.8, 5.9 &.10)
- The major divisions of the fishes alimentary canal are mouth, pharynx, esophagus,
stomach, intestine, rectum and related organs. These divisions are distinguishable
by the presence of valves or sphincters separating them. Associated with the canal
are two gland, the liver and the pancreas which deliver their secretion into intestinal
lumen via ducts. It is generally agreed that the morphology of the gastrointestinal
tracts in fishes is related to their feeding habits.
a) Esophagus:
- The great majority of fish have a short, wide esophagus with a mucus lubricated
entryway into the stomach. The esophagus mainly functions in the transport of
food into the stomach. Taste buds are common structures of the anterior part of the
esophagus wall of several teleosts. Several fishes with long slim body shapes
(anguillid eels) have long esophagus.
b) Stomach: (Fig. 5.11)
- The stomach of fish shows many variations in shape, but can be considered as organ
for short-term storage, mixing and primary digestion of food.
90
- Fish stomach show a great diversity and many attempts have made to
categorize them.
a) Elongated or J- shaped tube stomach, in many piscivores.
b) Sac or bag like stomach in most omnivores
c) Stomach modified into grinding organ . The stomach is reduced in overall size
but its walls, greatly thickened and muscularized.
d) Stomach absent in some omnivore, emphasizing plant sources of food.
c) Intestine:
- The intestine too, has many variations: (Fig. 5.12)

a) Carnivorous species tend to have shorter gut lengths.
b) Elongated and arranged in many folds in herbivorous species.
c) Intermediate gut length , corresponding to small zooplankton diet.
d) With a spiral valve or a large fold of absorptive tissue loosely wound
in a roll or folds and have improved efficiency of digestion and
absorption, Ex. Sharks and relatives.
- In the fish provided correlate the gut length to standard body length.
Ex. Valve 0.7 to 0.9 for carnivorores,
1.1 to 2.2 for omnivores, and
5.4 to 28.7 for herbivores.
d) Pyloric caecum: (Fig. 5.11)
- A pyloric caeca (blind tubes) can occur near the beginning of the midgut.
but are always absent in fish lacking stomach.
- The pyloric caeca and its associated mesenteries are thought to have a variety
of functions including digestion, storage or absorption.
- The histological structure and the enzyme content of the pyloric caeca appears
to be identical than those of the intestine.
91
(e) Digestive glands :
- The liver is a yellowish brown gland consisting of 2 main lobes that are sub-
divided into smaller lobes. One small lobe of the liver is isolated on each side
between the skin and the muscles.
- The gall bladder is a thin walled contractile sac between the intestine and the
right lobe of the liver, that stores the bile collected via the biliary ducts of the
liver. It functions as a temporary storage organ for liver secretion. Some deep
sea fishes have a vestigial gall bladder only and in others, the gall bladder is
completely absent.
- Bile duct is a short tube opening into the intestine, a little behind the pylorus.
- The pancreas is a well-developed diffused gland surrounding the blood vessels
between the lobes of the liver. A part of it extends over the surface of the
stomach also. There are smaller or larger nodules of pancreatic tissue in the
omentum, the suspensory tissues of the intestine, In addition to dispersion of
the pancreas into the liver to form a hepatopancreas among many spiny rayed
fishes.
- pyloric caeca, its secretions enter into the pyloric caeca directly. In other
cases, where the pancreas is inside the liver, it may empty directly into the gall
bladder.
92
(a) (b) (c)

Figure 5.1.A Variations in the development of the suctorial lips
and mouth in three types of sucker fishes
(a) shorthead (b) long head (c) long head modified
Figure 5.1.B Modifications in the types of mouth according to their shapes
93
Figure 5.2 Some structural adaptations of the mouth among fishes

(a) elephant fish (b) gar fish
(c) halfbeak (d) thick-lipped
(e) stargazer (f) longnose
(g) paddle fish
94
Figure 5.3 Some variations of teeth in fishes by structure and location
95
Figure 5.4 The pharyngeal mill of the cichlids fish.

Solid arrows show direction of movement of
pharyngeal bones; Broken arrows show food path
Figure 5.5 Three examples of pharyngeal bones from Cichlids

showing interspecific variation in form of
pharyngeal teeth
96
(a) (b) (c)
Figure 5.6 Relationship of gill rackers to feeding habits, showing

increasingly efficient sieve potential from left to right
(a) round white fish (b) lake white fish (c) blackfin cisco fish
97
Figure 5.7 Internal anatomical features of a spiny dogfish
98
Figure 5.8 Digestive system of shark
99
Figure 5.9 Internal anatomical features of a spiny-rayed bony fish
100
Figure 5.10 Internal anatomical features of a soft-rayed bony fish
101
Figure 5.11. Variations in shape, appendages, and lining of the anterior

portion of the digestive tract in three types of fishes.
102
Figure 5.12 Variations in intestinal length and other features

among carnivorous and herbivorous fishes
103
Exercise 6
104
AGE AND GROWTH DETERMINATION
The most frequently used method of age determination is the interpretation and counting
of growth zones or growth checks which appear on the hard parts of fishes. Those that
are considered to be formed annually are called year marks, annual marks, annual rings
or annuli. These formed during alternate periods of faster and slower growth (or no
growth at all), and reflect various environmental or internal influences. In temperate
regions the period of little or no growth usually occurs between the beginning of winter
and early summer. In general the greater the seasonal temperature differences, the clearer
are the annual marks. For this reason they are most distinct in northern and southern
parts of the world with a continental climate.
The choice of age determination methods:
Great care must be taken with any species for which no established ageing method
is available. The first step is to examine the scales and otoliths, and to prepare dried
bones, particularly flat ones such as the operculum, vertebrae and cross-sctions of spines
and fin rays, from three or four individuals. The aim is to find a structure that satisfies the
criteria.
The procedures used in age determination:
Definition and designation of age: The term ‘age-group’ expresses age in years, while
‘year-class’ refers to the fish produced in a given year. A consistent system for
designating age is necessary, regardless of the method by which age is determined.
Unfortunately there is not yet complete agreement on age designation or terminology.
By common usage the age of a fish is designated by reference to the annual marks on its
hard parts, and to some extent the season of year, rather than according to its exact length
of life (which in any event is usually not precisely known). A fish in its first growing
105
season belongs to age-group 0, and the successive stages may be called a larva (fry ocsac
fry) alevin (advanced fry), and fingerling (young-of-the-year or under-yearling).
A fish in its second growing season is said to be member of age-group 1, or simply age 1,
and is called a yearling. In the third growing season the fish is age 2 and is called a two
year-old, and so on.
An alternative numerical designation is age 1+ for yearlings, 2+ for two year olds etc.
The plus sign refers to the growth beyond the annual mark, which is appropriate enough
but not necessary.
TECHNIQUES FOR COLLECTING< CLEANING AND

PRESERVING STRUCTURES USED TO DETERMINE AGE
Scales:
In general there is no difficulty in taking scales from fishes. For many fishes a useful
instrument is a pair of pointed tweezers. When the scales are small, or embedded or
heavily coated in mucous, scales may be scraped from or dig out of the fish with a sharp
knife.
The most gently used area for scale sampling is the middle region of the side of the body.
The most useful scales for age assessment are usually large and symmetrical. For
comparability, the scales in any one study series should be taken from the same location
on all the fish, which may differ somewhat according to species.
Clean scales can sometimes be examined dry, but they tend to curl, so some kind of flat
mount is almost essential. Formerly scales were usually mounted on glass slides in a
glycerin-gelatin medium with the rough (outside) surface up.
A more recent and more satisfactory technique is to prepare an impression of the outer
surface of the scale. This is done by placing the scale, sculptured outer side down, on a
thin slide of plastic about 1 mm thick. The slide with the scale is place between two more
106
slides of plastic and is run through a roller press, using enough pressure to make a distinct
and complete impression. If large quantities of fish are being handled, scales from many
specimens can be mounted on large slides, and a heated vertical press is used to make the
impression. The slides with the sculptured impression are preserved for study. The use of
plastic slide impressions saves time, material and storage space and affords permanency.
Otoliths:
To obtain an otolith, a dissection of head bones is required. Of the three otolith on each
side, only the largest, the sagitta, is used. These should be removed with a pair of fine
tweezers to prevent breakage, and can be preserved dry in a paper envelopes or suitable
vials. Sometimes careful mechanical cleaning of an otolith is necessary, using lancet
needle. To brighten the otoliths and accentuate the appearance of growth rings, glycerin,
xylol, cedar wood oil, or cresol may be useful.
Bony parts:
The removal of bones usually means killing the specimen except where only fin spines or
rays are taken. The most frequently used bone are operculum, vertebrae, cleithum, fin
spines or spiny rays and soft rays.
After dissection, bones have to be cleaned of flesh and fat, and then dried. Opercula can
be removed quite quickly, using a scalpel to cut round the posterior edge (without
damaging the bone) and then twisting with the fingers. As soon as convenient after
removal they should be cleaned by soaking in hot or boiling water, fragments of flesh
remaining can be removed by a cloth or paper tissue held between the fingers and thumb.
The rings appear on drying and become clearer after storage. Thick parts of opercula can
be carefully filled or scraped to reveal inner layers and rings, or cleaned for projection by
immersion in xylol.
107
Age determination from scales:
The structures and appearance of the scales of bony fishes differ in different species.
Among higher bony fishes most of the upper surface of the scale is covered by ridges or
circuli. These are usually but not always more or less concentric, surrounding a central
spot or focus. In many species the ridges are interrupted by a series of radii that extend
from the focus to the margin, and may branch occasionally. The ridges represent the
margins of the successive thin plates of which the scale is constructed. The oral or
anterior field of the scale is usually the more useful one for identifying annual marks.
Two major scale types, cycloid and ctenoid, correspond in a general way to the division
teleosts into soft-rayed species with pelvic fins in the abdominal position, and species
having (usually) some spiny rays and with pelvic fins in the thoracic position. Ctenoid
scales usually differ from cycloid ones in having the posterior field variously adorned
with short conical teeth, which may interrupt the ridges and make them unrecognizable.
However in some species, or on some scales of a given species, the spines are reduced or
even completely missing. On ctenoid scales the circuli tend to be closer together than on
cycloid scales.
The following scale features have been useful, in various combinations, for identifying
annual marks in different fishes:
1. A zone of closely-spaced ridges is followed by a zone of widely-spread ridges; the
annulus is usually considered to be at the outer border of the closely-spaced ridges, but
sometimes in the central part or at the inner margin of such ridges.
2. A clear zone, devoid of ridges (perhaps with ridges absorbed), occurs between a zone
of closely-spaced ridges and a zone of widely-spaced ridges
3. Ridges become markedly discontinuous.
4. ‘Cutting over’ occurs, where one or two ridges appear to cut across several others. This
is usually discernible on the dorso-lateral and ventro-lateral parts of the scale.
5. In Clupeidae, where the ridges form two broad arcs the anterior field of the scale, the
annual mark is indicated by a slight bend or waviness in the ridges.
6. in some scales, especially ctenoid ones, the radii end, bend at the annual mark.
108
Age determination from otoliths:
Usually an otolith is examined from above by reflected light. It then exhibits opaque
zones of a white milky color and more translucent zones of a darker color. It is best to
place the otolith in a black dish and cover it with water, glycerin, xylol or cresol.
Alternatively, the otolith can be broken or cut transversely, and the broken surface
examined either dry or wetted with glycerin, with or without preliminary polishing.
Otoliths can also be prepared for identification by burning and breaking. On a burned
otolith, very distinct black and white rings appear at the surface of the fracture. The black
zones coincide with the boundary between opaque and transparent rings as seen in whole
mounts or sections. The carbonization method has been used successfully for the otoliths
of sole fish, carps, zander fishes and also the eel fishes.
Age determination from opercula and other bones:
On the opercula of fish from temperate waters, broader opaque zone correspond to the
more rapid growth of summer, and fade into narrow transparent winter rings which
usually end abruptly the start of the next summer zone. The rings show most clearly when
viewed by reflected light against any black background, which makes the summer zones
appear white and the winter zones black. The reverse is seen with transmitted light, with
which it may also be helpful to use polarized light. This is easiest done by rotating the
bone between two ‘crossed’ discs of Polaroid or in front of one piece of Polaroid while
wearing Polaroid glasses.
109
Figure 6.1 Scales with annual marks
110
Figure 6.2 Scales with annual marks from different locations
111
(a) (b)
Figure 6.3
(a) An example using an otolith of assigning a year class and /or birth
date, using information on the time of year in which it was collected
(b) A typical ctenoid scale, showing groups of concentric rings that can
be classified into annuli and interpreted as seasonal growth marks
112
Figure 6.4 Three methods of otolith extraction:

(a) Via the operculum above the gills
(b) By slicing the top of the skull off
(c) By a vertical skull cut in flatfishes
113
Figure 6.5 An otolith shows different zones and surfaces of growth rings
114
Figure 6.6 Drawing of a broken and burnt otolith

positioned in plasticine under a microscope
115
Figure 6.7 An operculum showing annuli
116
Figure 6.8 Annual growth marks on

(A) The vertebra, and
(B) The operculum of a bony fish
117
Figure 6.9 Diagram showing the two kinds of concentric

marks on the vertebral column of elasmobranchs
Figure 6.10 The shape of an elasmobranch vertebral centrum

and two planes of sectioning that are often useful
in aiding ring and band counting
118
Exercise 7
URINARY SYSTEM
Kidney structure and function:
The kidneys are seen as long reddish-brown structures running dorsally in the
body cavity above the gas bladder and beneath the vertebral column.
The functions of excretion and osmoregulation are closely related and are
performed by gills and kidneys in fishes. Although the gills are chiefly the respiratory
organs, they are also important as excretory and osmoregulatory organs. Kidneys play
the most important part in the excretion of nitrogenous wastes and in maintaining the
water-salt balance (homeostasis). A kidney consists of a large number of nephrons and
tubules which develop from infront backwards. The tubules of the anterior region
develop and become functional in early life and constitute the pronephros. Those of the
more posterior region develop and become functional in later life and form the
mesonephros. The ancestral craniate probably had a complete set of segmental tubules in
the trunk region forming an archinephros and opening into an archinephric duct. The
archinephros became differentiated into pro-, meso-, and meta-nephros. A functional
pronephros is found in fishes larvae and is later replaced by mesonephric tubules. These
open into the pronephric duct which, therefore, becomes the mesonephric duct. The
tubules of the more anterior region or mesonephros have a tendency to become reduced
and even degenerate, becoming converted into a lymphoid organ. However, the more
posterior tubules take up the excretory function and form the bulk of the adult kidney in
fishes. The posterior tubules behind the prone-phros, in fish and amphibia, are also
known as 'opisthonephros'
Kidney of Cyclostomes :
Hagfishes and the origin of the Glomerular kidney:
In hagfish, the kidney is simple and pronephric. The kidney structure provides
evidence that the ancestors of the hagfish inhabited freshwater.
Hagfishes kidney have glomeruli, but their blood pressure seems too low to force
filtration into the nephron. Glomerular filtrate does form and drains almost directly into
120
paired ducts. It is unclear whether the levels of some solutes are modified as the filtrate
moves along the ducts and is excreted.
Lamprey kidney:
Larvae of the lamprey ( develop in freshwater) have functional pronephric

kidneys until they are 12 to 15 millimeters long. The nephrostomes have funnels that
open into pericardial coelom. Coelomic fluids are swept into the nephrostome, at the site
of the glomerulus, the circulatory component of the pronephric tubule, substance to be
conserved and reabsorbed into the blood and the remaining aqueous filtrate passes by
duct to the urinogenital sinus.
In metamorphosis of the larvae, the mesonephric kidney takes over and the
pronephric openings into the coelom disappear. Arterial blood is filtered through the
glomerulus without selective retention of solutes, while the convoluted tubule is
instrumental in reabsorbing salts and other substance.
Kidney of Elasmobranchs:
Elasmobranchs have very complex kidney tubules, each being subdivided into
several distinct functional regions. Elamobranchs form a relatively large volume of renal
filtrate because their internal osmotic concentration allows them to absorb water even in
salty environments. Active transport of salts into and out of the tubule in the different
segments ensures that the fish excretes excess water and waste products while retaining
essential osmolytes, including the urea that it needs to maintain its high internal osmotic
concentration.
Kidney of Bony fishes
Teleosts have kidney tubules are less complex than those of elasmobranchs. In
freshwater teleosts, the glomeruli and different tubule segments are well developed and
produce a fairly large volume of dilute urine as a result of the constant influx of water
from the environment. Sodium and chloride are reabsorbed in the distal segment and
collecting duct, and little is lost in the urine.
121
Kidney tubules of many marine teleosts are considerably reduced compared to

freshwater teleosts. This results in a low rate of filtrate formation and decreased overall
urine output, there by helping prevent excess water loss.
Structure of the kidney:
The kidneys are paired, elongated structures placed above the alimentary canal
and are close to the vertebral column. The teleostean kidney is generally divided into
two portions, the head kidney and the trunk kidney, but in many species, these regions
can be distinguished by external examination. Generally, there are no conspicuous
differences in shape between the two sexes. According to Ogawa (1961), the marine
teleostean kidney can be divided into five types (Fig. 7.1)
Figure 7.1 Five types of teleostean kidneys.
1. The two kidneys are completely fused and there is no clear distinction
between trunk and head kidney, as in clupeidae.
2. The middle and posterior portions of the kidneys are fused. There is a clear
distinction between the trunk and head kidney, as in the marine catfishes
(Ploto-sidae) and eels (Anguillidae).
3. Only the posterior portions of the kidney are fused. The anterior portion is
represented by two slender branches. Head and trunk kidneys are clearly
distinguishable. Example : most marine fishes belonging to the Belonidae,
Scopelidae, Mugilidae, Scombridae, Carangidae, Cottidae and Pleuronectidae.
122
4. Only the extreme posterior portion of the kidney fused. There is no

distinction of head and trunk kidney, as in Syngnathidae (Sea horse and Pipe
fishes).
5. The two kidneys are completely separated as in Lophiidae.
Histology of the kidney:
The diagrammatic representation of a teleostean nephron (Fig. 7.2) shows the renal
corpuscle, neck region, different segments and the collecting duct.
Histologically, the trunk kidney is made up of a large number of nephrons, each

consisting of a renal corpuscle or the malphigian body of the tubule.
The intercellular space is full of lymphoidal tissue which is unevenly distributed (Fig.7.2)
The head kidney is generally made up of lymphoid, haematooietic, interregnal and

chromaffin (supra renal) tissue and is devoid of renal corpuscles and the tubules (Fig.7.2)
In some species, however, a few nephrons and collecting ducts may be present in the
lymphoid tissue of the head kidney, but the glomeruli and the renal corpuscles are usually
absent.
The head kidney is therefore, not excretory in function.
It is derived from the pronephric haematopoetic tissue but in some teleosts, the
mesonephros grows forwards extending into the head kidney.
The renal elements found in the head kidney of some teleosts are very likely
mesonephric.
123
Figure 7.2 Diagrammatic representation of a teleostean nephron, entire and in T.S.
124
Exercise 8
125
REPRODUCTIVE SYSTEM
INTRODUCTION:
Most fishes are egg-layers, but many bear living young. Live-bearing fishes
may be ovoviviparous, in which the eggs essentially simply hatch within the female,
or viviparous, in which the unborn young are supplied nourishment through the
mother’s tissues. In some ovoviviparous fishes the embryo develops in the egg while
the egg is still within its follicular covering within the ovary, and ovulation (or release
of the egg) and birth occurs at the same time. In other ovoviviparous forms the eggs
are released from the protective follicles in to the cavity of the hollow ovary, where
development continues. In some viviparous fishes the walls of the egg follicle are in
intimate contact with the embryo, supplying it with nourishment. In the viviparous
sharks, a part of the oviduct, or egg channel, is developed into a uterus, where the
modified yolk sacs of the young are closely joined to pockets within the uterus.
In live-bearing fishes and in some egg-layers, fertilization occurs internally,
and methods have been evolved for introducing the sperm into the females
body. In sharks the pelvic fins of the male are modified into intromittent organs called
myxopterigia, and in the male topminnows the anal fin modified into a similar –
functioning intromittent organ called the gonopodium.
At least three modes of reproduction- heterosexual, hermaphroditic, and
parthenogenesis—are found in fishes. In the most common form, heterosexual
reproduction, there are separate male and female parents, but even here there is
considerable variation. In some live-bearing fishes, the female is able to store sperm
for up to 8 or even 10 months, and this sperm is used to fertilize new batches of eggs
as they develop. In some cases, a female may carry sperm from several males at once.
In hermaphroditic reproduction, a single fish is both male and female,
produces both eggs and sperm (either at the same time or at different times), and
mates with other similar hermaphroditic fishes. External self fertilization occurs in
one hermaphroditic fish, which sheds egg and sperm simultaneously. In another,
internal self-fertilization may occur. In certain fishes there is a time sequence of
hermaphroditism, young fishes reversing their sex as they grow older.
126
In parthenogenetic reproduction, unfertilized eggs develop into embryos.

This is known to exist in at least one fish species, Poecilia formosa of the Amazon
River; however, even though development proceeds without fertilization in some of
these females, mating with a male is still required to stimulate egg development.
Parental care shows great diversity. Some fishes like the Atlantic herring;
form huge schools of males and females and freely shed their eggs and sperm (milt),
and abandon the eggs. Other fishes build nests and care for both the eggs and newly
hatched young. Others have evolved methods of carrying the eggs with them,
commonly in their mouths, but also in gill cavities or in special pouches on the body.
GENERAL ANATOMY:
Fish like all other vertebrates, reproduce sexually, and in the majority of
species, but not all, the eggs and sperm are formed in separate individuals. Whilst
simultaneous hermaphroditism and sex change do occur in an array of fish species,
the discussion of the reproductive system given here will be confined to general
description of the anatomy and developmental events occurring in dioecious,
gonochoristic species.
AGNATHA REPRODUCTIVE SYSTEM:
Both lampreys and hagfishes have unpaired gonad occupies most of the
coelomic cavity without a gonoduct. The eggs or sperm are shed into the body cavity
and then extruded through abdominal pores into the water.
Lampreys:
In the largest sea lampreys, (Petromyzon marinus), eggs counts have

ranged between 2400-236000, whilst in smaller non-parasitic forms may be 400-9000
eggs. Females of large anadromous lampreys produce 124,000 to 260,000 eggs. The
eggs are laid in ‘nests’, made in the stream bed, and hatch in a fortnight or so as small
pro-ammocoete larvae which are many respects ‘prototype’ chordates. These soon
127
change into the active ammocoete stages which burrow into silt bank and filter-feed
for several years. After five or more years, the ammocoetes metamorphose into adults.
Generally larval life is longer in parasitic species than in non-parasitic. Most lampreys
die after spawning, but some anadromous forms may survive to spawn second time.
Hagfishes:
Female lays few ellipsoidal, tough shell, leathery yolky eggs (20-30) and
2-3 cm long. The eggs have small hook-like structures on their ends for attaching to
the bottom and to each other. After 2 months or more the eggs hatch as small version
of the adult. Young are hermaphroditic at first, bearing both sets of sex organs later in
life, they will be either male or female but many change sex from season to season.
CARTILAGENOUS FISHES REPRODUCTIVE SYSTEM:
Elasmobranchs have paired penis or claspers formed from the inner

elements of the pelvic fin that roll up to form a tube with overlapping edges. These
male claspers transfer sperms to the female to fertilize the eggs in the oviduct, after
which they may either be retained or laid on the sea bed. In males, modified portions
of the kidney tubules and ducts also transport sperm and so can have either excretory
or reproductive functions. In females, however, the two organs are separated
structurally and functionally. Unmodified ducts drain female kidneys, and ovaries
release their eggs into oviducts lying near the posterior portion of the kidneys.
Male urinogenital system:
In males, the testes are paired, elongated oval structures at the front of body
cavity. Modified kidney tubules called efferent ductules transport sperm to the
archinephric duct. At the posterior end of each archinephric duct, seminal vesicles
connect to the sperm sac and contribute secretion to make up the seminal fluid, which
128
helps transport sperm. The sperm sac and seminal vesicle join to form the urinogenital
sinus, a cavity within urinogenital papilla.
Males have sacs that secrete fluid to lubricate medial groove in the clasper
to facilitate sperm transportation.
The kidney and gonads, collectively called the urinogenital system.

Although these two organs have entirely different functions, they are associated
morphologically and functionally.
Female urinogenital system:
In females, the ovaries are located at the anterior end of the body cavity
and suspended from the kidneys by the mesovarium membrane. The oviduct unite
anteriorly to form the ostium, the opening receiving the eggs , then they continue
backward to open into the cloaca on each side of the urinary papilla. The oviduct
includes an oviducal (nidamental or shell) gland, which secrets a shell around the
eggs. The oviducts end at the uterus, an enlarged portion housing the developing
embryo. In contrast to the male duct, the female archinephric duct is not convoluted
and is less prominent. Like males, females have a urinary papilla and a cloaca
receiving excretory and reproductive products, although in reproduction the female
cloaca serves as the birth canal. Eggs develop in the ovaries and, during ovulation,
burst through the ovarian walls into the coelom near the ostium. There the eggs begin
to descend into the oviducts, which are highly elastic. Eggs are fertilized in the
oviduct before the shell gland provides them a thin membranous covering, which later
breaks down and disappears. Embryos then gestate for a long time (around two years
for Squalus), nourished by the yolk suspended in a sac from the ventral side of the
embryo. Although this external yolk sac is usually used up fully at birth, a smaller
internal yolk sac remains within the newborn shark’s body cavity to provide more
sustenance. This process is not exactly the same in all species of elasmobranches. In
some forms, the females lay eggs containing a large amount of yolk encased in a
horny shell that then develop outside the mother. In other forms, the nutrition for the
129
developing embryos does not come from a yolk sac but rather is divided directly from
placental papillae (trophonemata) lining the uterine wall.
BONY FISHES REPRODUCTIVE SYSTEM:
The gonads occupy space in the coelom above the intestine and behind the
stomach.
Male reproductive system
The reproductive system of male bony fishes includes paired testes

located behind the stomach and duodenum, below the swim bladder, and just above
the intestine. The testes are lobulated and have longitudinal folds along their main
axis. Within these folds, the tubular vas deferens lead to exterior between the urinary
papilla and the anus. In all fishes except salmonoids, the vas deferens exit through the
body wall at the genital pore. They are not a part of the kidney as they are in
elasmobranchs.
Female reproductive system:
In females, the ovaries are usually paired eg sacs located in the same
position as the testes in the male. Some fish (e.g., the yellow pech) have a single
unpaired ovary. The oviducts, which except in salmonoids, are continuous with the
ovaries, receive either riped or fertilized eggs. Eggs are usually expelled through
abdominal pore. The system is variously modified to allow live- bearing in viviparous
females.
130
Figure 8.1 Urinogenital organs of a typical bony fish
131
Figure 8.2 Urinogenital organs of some common bony fishes
132
HISTOLOGY OF REPRODUCTIVE SYSTEM
Testes and spermatogenesis:
Spermatozoa, or sperm, are formed from the germ cells via a series of
developmental changes known collectively as spermatogenesis (Fig 8.3).
Spermatogenesis involves an initial proliferation of spermatogonia through repeated
mitotic divisions, and growth to form primary spermatocytes. Primary spermatocytes
then undergo reduction division (meiosis) to form secondary spermatocytes. Division
of the secondary spermatocytes results in the production of spermatids, which then
undergo metamorphosis to the motile spermatozoa.
The testicular structure of teleosts is somewhat variable, but two basic
types can be distinguished- the lobular and the tubular. In the testis of the lobular
type, which is seen in most teleosts, there are numerous lobules separated from each
other by thin layers of connective tissue (Fig. 8.3). The connective tissue which
extends from the testicular capsule effectively divides the testis into lobules, and it is
around the perimeter of the lobules that the germinal epithelium develops. This
epithelium consists of a layer of Sertoli cells with germ cells embedded within them.
During the course of spermatogenesis the primary spermatogonia
undergo numerous mitotic divisions, resulting in the production of cysts surrounded
by Sertoli cells. Each lobule of the testis contains numerous cysts. Within each cyst
the groups of germ cells divide in synchrony, so all of these cells are at the same stage
of development at any given time. However, within the lobules the spermatogenetic
process need not be synchronous, and germ cells at different stages of development
and differentiation may be found within a single lobule (Fig.8.3).
As spermatogenesis proceeds, the cysts increase in size and they
eventually rupture to release the spermatozoa into the lobular lumen. As progressively
more cysts reach maturity and rupture, the lumen becomes filed with spermatozoa.
Whilst it is usual that it is spermatozoa that are released from the cysts this is not
universal.
In some species, rupture of the cysts may occur at an earlier
developmental stage. Thus, spermatids, rather than spermatozoa, may be released
133
from the cysts. Differentiation and maturation of the spermatids to spermatozoa then
takes place within the lumen of the testicular lobule.
The lobular lumen is continuous with the sperm duct. Following a
variable period within the lumen, the sperm are released into the duct. This release of
sperm from the lobule into the sperm duct is known as spermiation.
The second type of testicular structure found in teleosts is restricted to
relatively few species, such as guppy, Poecilia reticulata. In the tubular testis the
primary spermatogonia are only found in cysts located at the blind end of the tubules
(Fig.8.3).
Figure 8.3 T.S of mature testes and seasonal changes in the testes.
134
Ovaries and oogenesis:
Ovarian follicle:
The ovary consists of germ cels, oogonia and oocytes, and their
surrounding follicle cells and supportive tissues. The ovarian follicles arise from the
germinal epithelium as each germ cell becomes surrounded by a layer of follicle cells
early in development.
With growth the follicle cells multiply and form a continuous layer
known as the granulosa cell layer. At the same time the outer layers of connective
tissues become organized to form a follicular envelope, or thecal cell layer. Thus, the
developing oocyte becomes surrounded by two major cell layers, the inner granulosa
layer and the outer thecal layer. The granuloa and thecal cell layers are separated from
each other by a distinct basement membrane (Fig.8.4)
Oogenesis:
The earliest stages of the development of the female germ cells are
similar to those seen in the initial stages of spermatogenesis (Fig.8.4). The germ cells
first undergo proliferation via a series of mitotic divisions and then they develop to
become primary oocytes. There is a period of growth, and a meiotic, reduction
division. The final reduction division, with expulsion of the second polar body, does
not occur until after fertilization, when the egg has been activated by the sperm.
The growth of the oocytes varies from species to species, with oocyte
enlargement being caused largely by the accumulation of yolk. Yolk accumulation
occurs during the developmental period known as vitellogenesis. The oocyte
enlargement may be considerable, with, for example, the oocytes of salmonoids
increasing in diameter from 20-50 μm to 4-5 mm during the course of development.
Oocyte development in teleosts:
Most teleosts have distinct breeding cycles, with spawning often
being limited to particular times of the year. Consequently, the ovaries may vary
greatly in appearance depending upon the stage of the breeding cycle.
For example, in matured or post-spawned females the ovaries and oviducts may
resemble thin strands, having a thread –or string –like appearance. On the other hand,
in the period immediately prior to spawning the ovaries may be so large and expanded
with oocytes that they almost completely fill the body cavity of the female.
135
The fact that the teleosts display a range of breeding cycles leads to
there being distinct paterns of oocyte development, and three broad categories have
been described. For example, all oocytes present within the ovary may be at the same
stage of development. This form of synchronous oocyte development is seen in
teleosts which spawn only once and then die, i.e., Semelparous species such as some
salmonoids and anguillids.
The majority of teleosts spawn more than once during the course of
their lives, i.e., are iteroparous. Amongst the iteroparous species two general patterns
of oocyte development have been identified.
The ovary may contain at least two distinct ‘populations’ of oocytes
at different developmental stages. Oocytes of a given population develop
synchronously. This type of developmental pattern is seen in the ovaries of teleosts
that spawn once per year within the space of a relatively short breeding season. Thus,
the ovary will always contain a reservoir of germ cells at an early stage of
development. The oocytes for successive annual spawnings are recruited from this
reservoir. The population or populations of oocytes recruited for spawning during the
course of a given breeding season will display synchronous, or almost synchronous,
development.
In some teleosts the ovaries may contain oocytes at all stages of
development, i.e., there is asynchronous oocyte development. This pattern is usually
seen in species which spawn many times during the course of a prolonged breeding
season.
136
Figure 8.4 T.S of mature ovary and seasonal changes in the ovaries.
137
Exercise 9
FECUNDITY MEASUREMENT
- Fecundity is the most common measure of reproductive potential in fishes, because

it is relatively easy to measure.
- It is the number of eggs in the ovaries of a female fish.
- In general, fecundity increases with the size of the female, with relationship.
F=aLb
where F = Fecundity,
L = Fish length, and
a and b are constant derived from the data (Bagenal 1978).
- The exponential relationship between fecundity and size in females seems to hold true
for many species (but, especially, for species that produce large numbers of eggs and
that spawn only once each year), but many factors complicate the interpretation of
fecundity data, especially in relation to the investment of energy. Some of these
complications are the relationship between fecundity and (1) fertility, (2) the
frequency of spawning, (3) parental care, (4) egg size, (5) environmental conditions,
and (6) population density (Bagenal 1978).
- By studying the relationships between fecundity (the number of eggs produced per
female) egg size and degree of parental care it is possible to make some generalizations
about reproductive patterns in fish (Tables 9.1 & 9.2). Thus, it can be said that the
following general principles appear to apply.
(1) Fecundity tends to be high where eggs and sperm are liberated freely into the water, is
less in those species giving some form of protection to the eggs and is lowest in species
that show parental care.
(2) Fecundity and egg size are inversely related, that is, fish produce many small eggs or a
few large ones.
(3) The production of pelagic eggs is largely restricted to marine species. Freshwater spcies
usually have non-buoyant, sticky eggs that are deposited on the substrate, or they build
some type of nest.
(4) Fecundity is higher in oviparous than in either ovoviviparous or viviparous species.
140
FECUNDITY DETERMINATION:
(1) Fish with the fecund ovary was carefully dissected, and ovary weighed and slitted
longitudinally and then turned inside-out before being immersed in Gilson's fluid for
fixing and preservation.
(2) Care should be taken not to lose any eggs by performing this operation in a whit's
tray, so that any eggs that was spilled fell in the tray to be put back into the specimen
jar.
Preservation of eggs:
Three different methods of preservation have been used:

(1) Gilson's Fluid: This fixative and preservative was first used in fecundity work by
Franz (1910) with European plaice. Simpson (1951) modified the recipe by doubling
the amount of acetic acid so that the mixture became:
100 ml 60% alcohol

880 ml water
15 ml 80% nitric acid
18 ml glacial acetic acid
20 g mercuric chloride
This modified Gilson's Fluid has been used by many investigators and has been found to
act satisfactorily with a wide range of species. Not only does the mixture harden the
eggs, but it also helps to liberate them and break down the ovarian tissue.
(2) Formalin: 4% or 5% formalin has been used by some workers (Scott 1962).
However in my experience formalin has not been satisfactory since the whole ovary
tends to become fixed in a hard mass and it is difficult to separate the eggs from
themselves and from the ovarian tissue.
141
(3) Boiling: Some early authors, for example Reibisch (1899) and Mitchell (1913), fixed
the ovaries by boiling them in water and then preserved them in formalin.
Details of Preservation:
If Gilson's Fluid is used, as recommended here, the treatment should proceed as follows.
- The ovaries, after they have been removed from the fish, should be split longitudinally
and turned inside out to assist penetration by the preservative.
- Large ovaries should be cut into separate portions.
- The ovaries should be preserved in labeled jars, shaken vigorously and left for at least
24 hours.
- Repeated shaking helps separate the eggs from the ovarian tissue and assists the
penetration of the preservative.
- The samples may be left for several months without disadvantage.
- If the eggs are not sufficiently hard to withstand handling by the time one wishes to
count them, gentle heat in an oven will hasten the process.
Washing:
- To clean the eggs, the Gilson's Fluid should be decanted and replaced by water.
- The periodic vigorous shaking will have liberated most of the eggs but the remainder
will have to be removed manually.
- The ovarian tissue free of eggs should be removed.
- The jar will now contain the free eggs in dilute Gilson's Fluid made cloudy
with minute fragments of ovarian tissue.
- By repeated filling with water and decanting the supernatant liquid the eggs will
become sufficiently washed.
- Simpson (1951) originally recommended washing with 70% alcohol to prevent
the eggs swelling and bursting, but water may be used providing the eggs have been
well preserved and are sufficiently hard.
142
Subsampling-Simpson's dry method:
- The preserved and washed eggs are poured into a filter paper in a funnel.
- When the liquid had drained away the paper and eggs are spread on blotting paper to
remove excess moisture.
- After about 20 minutes the eggs are spread on a filter paper tray made from sheets 30 x
45 cm or 45 x 60 cm, with the edges turned up.
- They are then left to dry until they can be moved without raising the surface of the
paper. They are left to air dry and moved about with the fingers to prevent clumps
forming. When they are thoroughly dry they can be stored in specimen tubes.
Analysis:
- Five approximately one-gram sub-sample counts were made for each ovary and the
means of these counts were used to determine the percentage number of yolky eggs per
gram sub-sample.
- The mean of these sub-samples was than used to estimate the fecundity of each female
by direct proportionality, using the following formula.
Es
F = Ws x Wg where,
F= fecundity
Es = mean number of eggs in sub-sample
Ws = mean weight of sub-sample (g)
Wg = total weight of Gonad (g)
- The resulting fecundity estimate were regressed on lengths and weights of parental
fish, and on ovary weight and egg size to provide information which can serve as
indication of changes in reproductive potential in the species.
143
Examples:
(1) Pampus argenteus (Euphrasen):

Fecundity estimates ranging from 28965-455661 (for 22 cm. & 28 cm. standard
Length respectively) (Dadzie, S et al.2000).
(2) Acanthopegarus latus:
A wide variation in fecundity was observed in this fish, even in fish of almost
the same size ranging from 368,000-3,837,000 ( Mean 1,277,062)
(Abou-Seedo et al.2003).
(3) Otolithes rubber:
Fecundity of this species is 77, 9,737 eggs.
144
Table 9.1 Relationships between fecundity, egg and larval size, and reproductive mode for a range of fish species
__________________________________________________________________________________________________________
Species Reproductive mode Fecundity Egg Size of larvae

_________________________ diameter ____________________________
(mm) (mm) (% adult length)
__________________________________________________________________________________________________________
Atlantic cod, Oviparous Marine 2 - 9 x 106 1.1-1.9 3.5 - 4.5 0.3

Gadus morhua pelagic
Plaice, Oviparous Marine 16 - 350 x 103 1.7-2.2 6–7 0.7
Pleuronectes platessa pelagic
Atlantic mackerel, Oviparous Marine 350-450 x 103 1.0-1.4 3–4 0.6
Scomber scombrus pelagic
Herring, Oviparous Marine 5 - 200 x 103 0.9-1.7 5–8 1.6
Clupea harengus demersal
Carp, Oviparous Freshwater 1802 - 530 x 103 0.9-1.6 5–6 0.6
Cyprinus carpio demersal
Atlantic salmon Oviparous Freshwater 1 - 10 x 103 5-6 15 – 25 1.3
Salmo salar demersal
Spurdog, Ovoviviparous 2 – 16 24-32 250 - 300 23
Squalus acanthias
Smoothbound, Viviparous 10 – 30 ~ 250 20
Mustelus mustelus
Guppy, Viviparous 10 – 50 6 – 10 13
Poecilia reticulate
_____________________________________________________________________________________________________________
145
Table 9.2 Fecundities of fish species displaying different levels of parental care
______________________________________________________________________________________________________
Egg type Degree of Fecundity Examples

parental care
_______________________________________________________________________________________________________
Pelagic, None 104 – 5 x 106 Atlantic cod, Gadus morhua

free-floating Turbot, Scophthalmus maximus
Atlantic mackerel, Scomber scombrus
Demersal, eggs None 103 – 105 Herring, Clupea harengus
stick to substrate Carp, Cyprinus carpio
or vegetation
Demersal Eggs buried in substrate, 5 x 103 – 5 x 104 Capelin, Mallotus villosus
hidden in holes or Salmonids
under rocks
Demersal Eggs guarded and cleaned 103 – 5 x 104 Lumpfish, Cyclopterus lumpus
(usually by male); eggs Blenny, Blennius pholis
may also be hidden Painted goby, Pomatoschistus pictus
Nest building; eggs 102 – 5 x 103 Threespine stickleback, Gasterosteus aculeatus
guarded and cleaned Bluegill sunfish, Lepomis macrochirus
(usually by male) Largemouth bass, Micropterus salmoides
Eggs carried by parent 50 – 1000 Sea horse, Hippocampus ramulosus
either in special brood Pipefish, Syngnathus acus
pouch or in the mouth Nile tilapia, Oreochromis niloticus
___________________________________________________________________________________________________________
146
IDENTIFICATION OF CARTILAGINOUS FISHES
Key to the Common Families of Cartilaginous Fishes (Sumich &

Dudley, 1996)
1 a. Pectoral fins not attached to sides of head, gill openings on sides ……………………………………sharks. 2
b. Pectoral fins attached to sides of head, gill openings under body ………………………... skates and rays. 14
2 a. Gills on sides, but otherwise skatelike …… ……. Squatinadae (angel sharks)
b. Body not skatelike ………………………………………………………...…………………………………. 3
3 a. Six or seven gill slits …………………………………………………………………………………………. 4

b. Five gill slits …………………………………………………………...…………………………………….. 5
4 a. First gill slit joined under throat …… …….. Chlamydoseachidae (frilled sharks)
b. First gill slit not joined under throat ….. ……. Hexanchidae (cow sharks)
5 a. Anal fin absent …………. …….. Squalidae (dogfish sharks)
b. Anal fin present …………………………………………….…………………………………………………. 6
6 a. Spine present at front edge of dorsal fins … .... Heterodontidae (horn sharks)
b. Spine absent at front edge of dorsal fins ………………………………..….………………………………... 7
7 a. Mouth at tip of snout, large white spots on body …….. ……. Rhincodonticae
(whale sharks)
b. Nose projects well forward of mouth …………………………………………………………...…………… 8
8 a. Caudal fin nearly as long as body ….. ….. Alopiidae (thresher sharks)
b. Caudal fin nearly as long as body …………………………………….………………………………………. 9
9 a. Eyes at ends of flattened head lobes ….. ….. Sphyrnidae (hammerhead sharks)
b. Head not as above …………..………………………………………...………...……………………………. 10
10 a. Front edge of first dorsal fin behind ….. ….. Scyliorhinidae (cat sharks)
front edge of pelvic fins
147
b. Front edge of first dorsal fin well in front of pelvic fins ………………………………..…….……………. 11
11 a. Upper and lower lobes of caudal ………. …... Lamnidae (mackerel sharks)
fin nearly equal
b. Upper lobe of caudal fin much larger than lower lobe ……………………………………………………. 12
12 a. Gill silts very long, extending … ….. Cetorhinidae (basking sharks)

to ventral midline
b. Gill silts not extending under body …………………………...………………………………………....... 13
13 a. Front edge of pectoral fins ……… …… Odontaspididae (sand sharks)

behind fifth gill slit
b. Front edge of pectoral fins under …….. …Carcharhinnidae (reguiem sharks)

fourth or fifth fill slit ……………………………………………………...………………………………..........
14 a. Dorsal fins absent ……………………………………………...………………………………………….. 15

b. Dorsal fins absent ………………………………………………...……………………………………….. 16
15 a. Body wider than long, very short tail …… ….. Gymnuridae (butterfly rays)
b. Body about as wide as long, ……… ……. Dasyatididae (stingrays)

longer tail with venomous spine
16 a. One dorsal fin ……………………………………………………………………………………………... 17

b. Two dorsal fins ……………………………………………………………………………………………. 18
17 a. Large lateral lobe on each side of head ……… ……. Mobulidae (manta rays)
b. No lateral lobe on side of head ……. ……… Myliobarididae (eagle rays)
148
18 a. Caudal fin absent ……… ……. Rajidae (skates)
b. Caudal fin present …………………………………………………………………………………….…… 19
19 a. Body smooth, no spines on back …… ….……. Torpedinidae (electric ray)
b. Body sausage-shaped, without spines ………………………………………………………………...…... 20
20 a. Body with three rows of spines on dorsal side ……… ……. Rhinobatidae
(guitarfishes)
b. Body with a single row of spines on dorsal side ……. …… Platyrhinididae

(thornbacks)
149
IDENTIFICATION OF BONY FISHES
Key to the More Common Families of Bony Fishes: (Sumich &

Dudley, 1996)
1 a. Eyes on same side of head …………………………………………………………………………………… 2

b. Eyes on same side of head …………………………………………………………………………………… 3
2 a. Caudal fin absent …… .……. Cynoglossidae (tonguefishes)
b. Caudal fin present …… …….. Bothid (halibuts and soles) ;

Pleuronectidae (turbots, soles, and flounders)
3 a. Pelvic fins absent …………………………………………………………………………………………..… 4

b. Pelvic fins present ………………………………………………………………...……………………...… 15
4 a. Body not eellike ……………………………………………………………………………………………… 5

b. Body eellike ………………………………………………………………………………………………… 10
5 a. Caudal fin absent, body rounded …… ……. Molidae (sunfishes)
b. Caudal fin present ………………………………………………………………………….………………… 6
6 a. Caudal fin very tall and rigid, upper jaw …… ……Xiphidae (swordfishes)
flattened and swordlike
b. Caudal fin soft, about as tall as broad ………………………………………………...……………...……… 7
7 a. Dorsal fin with soft rays and spines ……………………………………………………………………….… 8

b. Dorsal fin with soft rays only ….……………………………………………………………………….….… 9
8 a. Dorsal fin spines separated from soft rays …… …. Balistidae (triggefishes)
b. Dorsal fin spines connected to fin with soft rays … …. Stromateidae (butterfishes)
9 a. Upper teeth fused into one, skin spiny …… …. Diodontidae (porcupine fishes)
b. Upper teeth fused into two, skin smooth … …. Tetraodontidae (puffers)
150
10 a. Pectoral fins absent …… ……. Muraenidae (moray eels)
b. Pectoral fins present ……………………………………………………………………….……………… 11
11 a. Dorsal fin without spines …… ……. Ophichthidae (snake eels)
b. Dorsal fin with spines ……………………………………………………………………….…………….. 12
12 a. Body encased in bony plates …… ……. Syngnathidae (sea horses and pipefishes)
b. Body not encased in bony plates ……………………………………………………………………...…... 13
13 a. Caudal fin forked …… ……. Ammodytidae (sand lances)
b. Caudal fin not forked …………………………………………………………………………………….... 14
14 a. Anterior portion of dorsal fin spiny … ……. Cebidichthyidae (monkey-faced eels)
b. Dorsal fin entirely spiny …… ..... Pholididae (gunnels)
15 a. Pelvic fins abdominal ………………………………………………………………………………...…… 16

b. Pelvic fins thoracic ….………..………………………………………….……………………...………… 31
16 a. One dorsal fin ……………………………………………………………………………...……………… 17

b. Two dorsal fins ….………………………………………………………………………………………… 23
17 a. Caudal and dorsal fins rounded … ……. Cyprinodontidae (killifishes)
b. Caudal and dorsal fins not rounded ………………………………………….……………………………….. 18
18 a. Lateral lines absent ……………………………………………………………………...………………… 19

b. Lateral lines present …………………………………………………………………...………………..… 21
19 a. Photophores present … ... Gonostomatidae (bristlemouths)
b. Photophores present ………………………………………………………...…………………………….. 20
20 a. Tips of jaw and snout even … ……. Clupeidae (herrings and sardines)
b. Tips of snout projects over lower jaw …… ..... Engraulidae (anchovies)
21 a. Five to seven finlets following dorsal and anal fins … ..... Scomberesocidae
(sauries)
151
b. Finlets absent …………………………………………………………..………………………………….. 22
22 a. Pectoral fin longer than one-third of body … …. Exocoelidae (flying fishes)
b. Pectoral fin shorter than one-third of body …… ..... Belonidae (needle fishes)
23 a. Both dorsal fins with rays …………………………………………………………………….…………… 24
b. Second dorsal fin soft, without rays …………………………………….………………………………… 27
24 a. Snout tubular ….… ….. Centriscidae (snipefishes)
b. Snout not tubular ……………………………………………………………………...…………………... 25
25 a. Lower jaw projecting, teeth large …… ….. Sphyraenidae (barracudas)
b. Jaws even, teeth small or absent ………………………………………………………….……………….. 26
26 a. Three spines in anal fin …… ……. Mugilidae (mullets)
b. One spine in anal fin …… ...... Atherinidae (silversides)
27 a. Origin of dorsal fin opposite or posterior to pelvic fin origin ……………………………………...…...… 28
b. Origin of dorsal fin anterior to origin of pelvic fin ………….………………………………...………..… 29
28 a. Lower teeth large, caninelike .. …. Synodontidae (lizard fishes)
b. Lower teeth small, few …… ..... Osmeridae (smelts)
29 a. Photophores absent …… ….. Salmonidae (salmon)
b. Photophores present ………………………………………………………………...…………………….. 30
30 a. Body strongly compressed …. …... Sternoptychidae (hatchetfishes)
b. Body not strongly compressed …… ..... Myctophidae (lantern fishes)
31 a. Pelvic fins modified as a cone or disc ……………………......................................................................… 32
152
b. Pelvic fins not so modified ………….………………………………………………………………….… 33
32 a. One dorsal fin ……… …... Gobiesocidae (clingfishes)
b. Two dorsal fins ………… ..... Gobiidae (gobies)
33 a. Pelvic fins with one spine and five soft rays ……………………………………………...…………….… 34
b. Pelvic fins without soft rays ………….………………………………………………...……………….… 51
34 a. Gill openings below or behind pectoral fin … ….. Antennariidae (frogfishes)
b. Gill openings anterior to pectoral fins …………………………………………………………………….. 35
35 a. First dorsal fin modified as a …… ….. Echeneididae (remoras)
b. First dorsal fin not so modified ………………………………………………………………………..….. 36
36 a. Dorsal fin followed by …… ….. Scombridae (tunas and mackerels)
b. Dorsal fin followed by one or no finlets …………………………………………………….…………….. 37
37 a. Dorsal fin spines absent …… ….. Coryphaenidae (dolphinfishes)
b. Spines present in dorsal fin ……………………………………………………………………………….. 38
38 a. Lowermost rays of pectoral fins … ….. Triglidae (sea robins)
b. Lowermost rays of pectoral fins not separate ………………………………………………….………….. 39
39 a. Anal fin spines absent …… ….. Cottidae (sculpins)
b. Anal fin spines present ………………………………………………………………………...………….. 40
40 a. Dorsal fins separated …… ….. Anaplopomatidae (sablefishes)
b. Dorsal fins not separated ………………………………………………………………………………….. 41
153
41 a. Five spines on preopercle …… ….. Scorpaenidae (rockfishes)
b. Preopercle without five spines ………………………………………………………...………………….. 42
42 a. Suborbital stay (bone below eye) present …… ….. Hexigrammidae

(greenlings)
b. Suborbital stay absent ……………………………………………………………………………………... 43
43 a. Anal fin with one spine or two spines …………………………………………………...……………...… 44
b. Anal fin with three or four spines ….……………………………………………………………………… 46
44 a. Lateral line extending onto caudal fin …… ….. Scianidae (croakers)
b. Lateral line not extending onto caudal fin ………………………………………………………………… 45
45 a. Dorsal fin same height throughout ……… …... Branchiostegidae

(ocean whitefishes)
b. Second dorsal fin higher than first …… ..... Pomacentridae (blacksmiths)
46 a. Two anal fin spines isolated from remainder of fin … ….. Carangidae (jacks)
b. Anal fin spines connected to remainder of fin ………………………………………………………...….. 47
47 a. Snout pointed, teeth very small …… ….. Chaetodontidae (butterfly fishes)
b. Snout not pointed ………………………………………………………………………………...……….. 48
48 a. More than ten anal fin soft rays … ….. Embiotocidae (surfperches)
b. Ten or fewer anal fin soft rays …………………………………………………………………………….. 49
49 a. Anterior teeth caninelike … ….. Labridae (wrasses)
b. Anterior teeth not caninelike ……………………………………………………………...………………. 50
154
50 a. Soft-rayed portion of anal fin shorter than soft-rayed

portion of dorsal fin ……… …... Serranidae (sea basses and groupers)
b. Soft-rayed portions of anal and …….. …... Kyphosidae (sea chubs)

dorsal fins about equal ….
51 a. Pelvic fin with less than five soft rays…………………………………………………………………...… 52
b. Pelvic fin with more than five soft rays …………………………………………………...……………… 64
52 a. Upper jaw projected into a spear … ….. Istiophoridae

(sailfishes and marlins)
b. Upper jaw not as a spear …………………………………………………………………………………... 53
53 a. Photophores present … ….. Batrachoididae (midshipment)
b. Photophores absent ………………………………………………………………………………………... 54
54 a. Body encased in bony plates … ….. Agonidae (poachers)
b. Body not encased in bony plates …………………………………………………….……………………. 55
55 a. Isolated spines anterior to dorsal fin … ….. Gasterosteidae (sticklebacks)
b. No isolated spines anterior to dorsal fin ……………………………………………………………………56
56 a. Body not elongate, eelike ………………………………………………………………….....................… 57
b. Body elongate, eelike ….…………………………………………………………………...………...…… 59
57 a. Anal fin spines absent ……….. ….. Cottidae (sculpins)
b. Anal fin spines present ………………………………………………………………...………………..... 58
58 a. More spines than soft rays in dorsal fin ……… …... Clinidae
(fringeheads and kelpfishes)
b. Fewer spines than soft rays in dorsal fin …… ..... Blenniidae (blennies)
155
59 a. Dorsal fin entirely of spines ……………………………………………………………….....................… 60
b. Dorsal fin entirely of soft rays ….……………………………………………………………………….… 61
60 a. Anus positioned in anterior half of body ……… …...Stichaeidae

(picklebacks)
b. Anus positioned in posterior half of body ……… …...Pholididae (gunnels)
61 a. Tail forked ……….. ….. Trichiuridae (cutlass fishes)
b. Tail tapered, not forked ………………………………………………………………...……………….... 62
62 a. Pelvic fins on chin or under eyes ……….. ….. Ophidiidae (cusk eels)
b. Pelvic fins not on chin or under eyes ………………………………………………………………........... 63
63 a. Pelvic fins clublike ……… …... Zoarcidae (eelpouts)
b. Pelvic fins long, not clublike ……… …... Brotulidae (brotulas)
64 a. No barbels under chin …… ….. Merlucciidae (hakes)
b. Barbels present ……………………………………………………………...………………….................. 65
65 a. Caudal fin absent, tail pointed …… ….. Macrouridae (grenadiers)
b. Caudal fin present ……………………………………………………………............................................. 66
66 a. Three dorsal fins ……… …..…... Gadidae (cods)
b. Two dorsal fins ……… ..... Moridae (codlings)
156
REFERENCES
Romer, A.S. & Parsons, T.S. 1977. The Vertebrate Body. 5th ed. Saunders, Philadelphia.
(6th ed. 1985)
Nelson, J. S. 2006. Fishes of the World. John Wiley & Sons, Inc…
Sumich, J.L & Dudley, G. H. 1996. Laboratory & Field Investigations in Marine Life
(6th ed.), Gulf Coast Version
Lagler, K.F., Bardach, J.E.,Miller, R.R. & Passino, D.R.M. 1977. Ichthyology (2nd ed.),
John Wiley & Sons.
Khanna, S.S. An introduction to fishes. 2003. Central Book Depot, Allahabad
157
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Preface & Acknowledgements I

1. General characteristics, Taxonomy and Agnatha (Jawless fishes) 1

2. Gnathostomata (Jaw fishes) 27

3. Skin, Skin derivatives & Fins 42

4. Blood & its Circulation 57

6. Age & Growth determination 104

7. Urinary system 119

8. Reproductive system 125

9. Fecundity measurement 138

10. Identification Key 147

11. References 157

A number of species possess barbells which are excellent organs of touch.

 Class Petromyzontida or Hyperoartia

AGNATHA (Jawless Fish)

- The cyclostomes (Fig.1.2) are characterized by the presence of a suctorial mouth

Examples: Myxine glutinosa, Bdellostoma (Eptatretus) cirrhatum, & Paramyxine etc.

GNATHOSTOMATA (Jaw Fishes)

Class Chondrichthyes divided into 2 subclasses.

Sub-class: Elasmobranchii: (Figs. 1.4 &1.5)

Ex: Frill sharks,

Sub-class: Holocephali: (Fig. 1.6)

Ex: Plownose chimaeras,

- The skull is usually hyostylie (or) 'amphistylic'.

The Class Osteichthyes divided into 2 sub-classes, Actinopterygii and Crossopterygii.

II Sub-class: Sarcopterygii or Crossopterygii ( Lobe fin fishes): Eg: Latimeria sp

Infraclass : Dipnoi : (Fig.1.9)

AGNATHA (JAWLESS FISHES)

- Petromyzon has an elongated eel-like, nearly cylindrical body with a laterally

- The skin of lamprey is slimy and is composed of several layers of epidermis.

- In addition to these, hag-fishes have segmentally arranged 'thread-cells' forming

- In lampreys the buccal funnel is suctorial and armed with teeth.

- The respiratory organs consist of a number of branchial sacs with numerous

- The brain of Petromyzon is in a relatively primitive condition.

- The kidneys are very small in the hag-fishes.

Figure 1.1 Ostracoderms

Figure 1.2 Cyclostomes

Figure 1.3 Placoderms (Extinct)

Figure 1.4 Elasmobranchii

SKATE OR GUITAR FISH

HAMMER HEADED SHARK

Figure 1.5 Elasmobranchii contd……

Figure 1.6 Holocephali

ACIPENSER (STURGEON FISH)

AMIA (BOWFIN FISH)

ATRACTOSTEUS (GAR FISH)

Figure 1.7 Actinopterygii

Figure 1.8 Sarcopterygii

Figure 1.9 Dipnoi

Figure 1.11 L.S. of Head region of Petromyzon

Figure 1.12 L.S. of Head region of Myxine

Structure and Body Shape:

- The great ecological diversity of fishes is reflected in the astonishing variety of

- The four groups of cartilaginous fishes (Chondrichthyes) have several distinguishing

- Commonly the fish body is torpedo-shaped (fusiform).

2. Body covering (Skin):

- An ordinary fish is covered with a relatively tough skin.

Figure 2.1 Some meristic and morphometric characters