Fish Systematics

Fish systematics is the formal description and organization of fish taxa into systems. The term
"fish" describes any non-tetrapod chordate, (i.e., an animal with a backbone), that has gills
throughout life and has limbs, if any, in the shape of fins. Unlike groupings such as birds or
mammals, fish are not a single clade but a paraphyletic collection of taxa, including jawless,
cartilaginous and skeletal types. Systematics is the study of the diversity of organisms and the
relationships between these organisms. Its goal is to classify species and detect phylogenies – a
purpose that differs from the objectives of taxonomy. Classification consists of recognizing and
defining groups or taxa (that is, a set of organisms that share a specific character) that
taxonomists will then have to name. According to the hierarchical classification proposed by
Linnaeus (1758), each level of the hierarchy corresponds to a taxon name. While ideas governing
classification in particular have evolved significantly since the 18th century, the basic rules
stated by Linnaeus remain solid. The discipline currently known as biosystematics is a modern
approach to taxonomy and phylogeny that makes use of information from different sources:
morphology, genetics, biology, parasite specificity, behaviour, and ecology.
Agnatha
Jawless fish
Jawless fish were the earliest fish to evolve. They have no jaw, no scales, no paired fins, and no
bony skeleton. Their skin is smooth and soft to the touch, and they are very flexible. Instead of
a jaw, they possess an oral sucker. They use this to fasten onto other fish, and then use their
rasp-like teeth to grind through their host's skin into the viscera. Jawless fish inhabit both fresh
and salt water environments. Some are anadromous, moving between both fresh and salt
water.

Agnathan, (superclass Agnatha), any member of the group of primitive jawless fishes that
includes the lampreys (order Petromyzoniformes), hagfishes (order Myxiniformes), and several
extinct groups.

Hagfishes are minor pests of commercial food fisheries of the North Atlantic, but lampreys,
because of their parasitic habit, have been a serious pest of food fisheries in the Great Lakes in
North America, where they have reduced the numbers of lake trout and other species.
Agnathans are otherwise of little economic importance. The group is of great evolutionary
interest, however, because it includes the oldest known craniate fossils and because the living
agnathans have many primitive characteristics.

General features

The body of the hagfish is soft-skinned, scaleless, and nearly cylindrical, with a single nostril at
the anterior end, overlying the mouth, and a low caudal fin around the tail. In some species the
gills open to the surface through separate pores; however, in others the gills open to a common
duct, which in turn opens to the surface through a single pore. The eyes are vestigial and
covered by skin. All of the approximately 70 known species are restricted to cold, marine
bottom waters at depths ranging from 10 metres (about 33 feet) in high latitudes to 1,300
metres (about 4,300 feet) in equatorial oceans. Adults are 40 to 80 cm (about 15 to 30 inches)
long. All species are superficially similar except in the number and position of the gill apertures.

Lampreys, which number about 43 species, are found in cool, fresh, and coastal waters of all
continents except Africa. All species are rather similar. The body is smooth, scaleless, and eel-
shaped, with well-developed dorsal and caudal fins; the mouth is surrounded by a suctorial oral
disk bearing horny teeth. The eyes are well developed, and the single nostril is on the top of the
head. Lampreys possess seven pairs of external gills. Adults range from 15 to 100 cm (6 to 39
inches) long. Although the gonad of a hagfish usually includes both ovary and testis, there is no
evidence of either hermaphroditism (the reproductive organs of both sexes functioning in the
same individual) or self-fertilization. A female produces a small number of tough-skinned yolk-
filled eggs about 2 cm (0.8 inch) long that hatch into miniature adults.

Hagfishes locate their food by scent. Although some are known to eat fishes immobilized in
nets, the best-studied species, Myxine glutinosa, normally feeds on soft-bodied invertebrates
and larger dead animals. Myxine burrows into soft marine sediments and rests with only the tip
of the head protruding. During respiration, water enters through the nostril and passes by a
nasopharyngeal duct to the pharynx and gills. When stimulated by the scent of a dead fish,
Myxine leaves its burrow and swims against the current. On making contact with the fish,
Myxine coils around it and bites into it by protruding and retracting the comblike horny tooth
plates on the floor of the mouth. Along each side of Myxine’s body is a row of prominent glands
that produce a gelatinous slime when it is disturbed.

Lampreys mate in a nest like depression excavated by the male in the gravel bed of a stream;
the numerous eggs, about 1 mm (0.04 inch) in diameter, lodge in the gravel around the nest.
The egg hatches into an ammocoete larva—a blind, wormlike animal that burrows in silt. The
larva’s mouth is overhung by a hoodlike upper lip that protrudes above the surface of the silt. A
continuous stream of water passes in through the mouth and out through the seven pairs of
gills. Microscopic plants, the food of the ammocoete, are filtered from this respiratory current
by strands of mucus produced by the endostyle, which is a gland in the floor of the pharynx.
Cartilaginous fish
Cartilaginous fish have a cartilaginous skeleton. However, their ancestors were bony animals,
and were the first fish to develop paired fins. Cartilaginous fish don't have swim bladders. Their
skin is covered in placoid scales (dermal denticles) that are as rough as sandpaper. Cartilaginous
fish do not have bone marrow, the spleen and special tissue around the gonads produces red
blood cells. Their tails can be asymmetric, with the upper lobe longer than the lower lobe. Some
cartilaginous fishes possess an organ called a Leydig's organ which also produces red blood
cells. There are over 980 species of cartilaginous fish. They include sharks, rays and chimaera.

Bony fish
Bony fish include the lobe-finned fish and the ray finned fish. The lobe-finned fish is the class of
fleshy finned fishes, consisting of lungfish and coelacanths. They are bony fish with fleshy, lobed
paired fins, which are joined to the body by a single bone. These fins evolved into the legs of
the first tetrapod land vertebrates, amphibians. Ray finned fishes are so-called because they
possess lepidotrichia or "fin rays", their fins being webs of skin supported by bony or horny
spines ("rays").

There are three types of ray finned fishes: the chondrosteans, holosteans, and teleosts. The
chondrosteans and holosteans are among the earlier fish to evolve, and share characteristics
with both teleosts and sharks. In comparison with the other chondrosteans, the holosteans are
closer to the teleosts and further from sharks.

Teleosts
Teleosts are the most advanced or "modern" fishes. They are overwhelmingly the dominant
class of fishes with nearly 30,000 species, covering about 96 percent of all extant fish species.
They are ubiquitous throughout fresh water and marine environments from the deep sea to the
highest mountain streams. Included are nearly all the important commercial and recreational
fishes.Teleosts have a movable maxilla and premaxilla and corresponding modifications in the
jaw musculature. These modifications make it possible for teleosts to protrude their jaws
outwards from the mouth. The caudal fin is homocercal, meaning the upper and lower lobes
are about equal in size. The spine ends at the caudal peduncle, distinguishing this group from
those in which the spine extends into the upper lobe of the caudal fin.

General principles of taxonomy

The rules of classification Classification of the living world is hierarchical, with groups included
in larger sets that do not overlap. But this hierarchy can be based on different principles.
Phenetic hierarchy is based on the similarity of appearance of the classified groups: number
and position of fins, presence or absence of barbels, number of fin rays, etc. Phylogenetic
hierarchy, meanwhile, is based on evolutionary relationship: groups are defined according to
the closeness of their relationships and the age of their common ancestors. These two
classification systems, which gave rise to different schools of thought, can provide concordant
results but may also differ in their conclusions. The numerical taxonomy school fosters phenetic
classification, whereas phylogenetic classification is supported by the so-called cladistic school.
Supporters of numerical taxonomy believe that organisms sharing common characteristics have
a similar evolutionary history, but do not make hypotheses on genealogy. By using a large
number of characters in a large number of individuals, and assigning them equal weights,
statistical methods are expected to identify homogeneous sets. However, morphological
convergences during evolution occasionally led to artificial groupings, and the biases that could
be generated by statistical methods were also criticized (Ridley, 1989). Cladistic classification
(sometimes called Hennigean classification) is based on the principle that during evolution, an
ancestral species gave rise to two daughter species. For each species, it is therefore necessary
to determine with which other species it shares the most recent common ancestor, because
this will form the basis for the first group. A monophyletic group is derived from a single
common ancestor, whereas a polyphyletic group includes species that have similarities but are
not direct descendants of a common ancestor. Unlike phenetic relationships, phylogenetic
relationships cannot be observed directly. How, then, can they be identified? The method
proposed by Hennig (1950) consists of looking for characters that can be qualified as
evolutionary innovations. We can thus distinguish evolved or apomorph characters in
opposition to ancestral characters, referred to as primitive or plesiomorph. This method, which
often used comparisons with distant groups to determine if characters were evolved or
primitive, also shows a bias – it considers that evolution always proceeds from a plesiomorph
state to an apomorph state, and never the reverse (Daget, 1980).
Trends in classification

The first fossil vertebrates were found in the rocks of ordovician period and named as
Ostracoderms. They are small, jawless, bony, fish like form related to cyclostomes. Devonian
period is known as “The Golden Age of Fishes”. Swedish naturalist, Peter Artedi (1705-1735) is
known as “Father of Ichthyology”. Artedi established the methods and principles of systematic
Ichthyology so perfectly even Linnaeus could not alter it. Cuvier was the first to frame the
classification on the basis of external and internal characters of fishes. The classification
proposed by Muller, Jordan, Regan, Berg, Romer and Nelson considered to be the recent
classification of bony fishes. Currently, Nelson (1994) classification of fishes is in practice.

Muller classified fishes into six (6) sub-classes Class: Pisces Subclasses: Six 1. Dipnoi (Lung fishes)
2. Teleostei (Bony fishes) 3. Ganoidei (Polypterus, Amia, Lepidos) 4. Elasmobranchi (Sharks,
Rays, & Rat fishes) 5. Marsipobranchii (Cyclostomes) 6. Leptocardii (Amphioxux).
L. S. Berg’s (1940) classified fishes into seven (7) classes Series: Pisces Class: Seven 1.
Pterichthys 2. Coccostei 3. Acanthodii 4. Elasmobranchii 5. Holocephali 6. Dipnoi 7. Teleostomi.
The teleostomii is further divided into crossopterygii and actinopterygii.

Romer’s (1971) A. S. Romer (1971) arranged fishes into four classes under super-class pisces.
Super-class: Pisces Class: Four 1. Agnatha (Jawless vertebrates) 2. Placodermii (armoured or
plate skinned fishes) 3. Chondrichthyes (Cartilagenous fishes) 4. Osteichthyes (Bony Fishes)

Nelson’s Classification (1994) J. S. Nelson (1994) presented the classification of fishes in general
in a linear order which reflects their postulated evolutionary relationship (cladistic approcah).
This includes Subphylum: Vertebrates Super-classes: 1. Agnatha 2. Gnathostomata. The
agnatha possess two classes 1. Cephalaspidomorphii 2. Pteraspidomorphii. The superclass
gnathostomata divided into two grades 1. Pisces 2.Tetrapoda

Grade: Pisces Subgrade Elasmobranchomorphii and Teleostomi. The elamobranchomorphii

includes the classes Class 1. Placodermii 2. Chondrichthyes which is further divided into 1.
Elasmobranchii 2. Holocepahali. The teleostomii includes class Class 1. Acanthodii (extinct) 2.
Osteichthyes. The osteichthyes is divided into subclass 1. Dipeustei 2. Crossopterygii 3.
Brachiopterrygii 4. Actinopterygii. The actinopterygii includes the Infraclasses 1. Chontrostei 2.
Holostei 3. Halecostomi (extinct) 4. Teleostei

Classification of fishes with reference to Nepal

Fishes have been classified in different ways from time to time. The earliest classification was
proposed by J. Muller (1844) who proposed seven sub-classes under the class ‘Pisces’. Later,
Berg (1940) and Romer (1959) have given detailed classification of fishes, which have generally
been accepted worldwide. In this book, we follow the classification given by Berg (1940), who
divided the Super-class (Series) Pisces into seven classes: (1) Acanthodii, (2) Coccostei, (3)
Pterichthys, (4) Elasmobranchii, (5) Holocephali, (6) Dipnoi, and (7) Teleostomi (Figure 2.1). Of
these, the first three classes are completely extinct and are collectively known as Placodermi
(meaning plate-skinned). Thus, the modern fishes are represented only by four major classes,
i.e., Elasmobranchii, Holocephali, Dipnoi and Teleostomi. Out of these, the fishes of Nepal
belong to the sub-class Actinopterygii of the class Teleostomi. Berg has divided the sub-class
Actinopterygii into a series of 59 orders. Out of these, only 11 orders are represented in
freshwater ecosystem of Nepal.

The total number of fish species represented in Nepal has been controvercial and is a subject of
study. Different fish biologists have reported different number of fish species present in Nepal,
which ranged from 228-232. According to Shrestha (2008), a total of 232 species belonging to
114 genera under 36 families and 11 orders existing in culture or natural water bodies of Nepal.
Recently, Shrestha and Pandit (2012) have described one more exotic species, Pangasius
hypophthalmus (Sauvage), which has been commercially cultured in the terai region of Nepal
since few years. This indicated that the total number of fish species present in Nepal might be
more than 232.

List of orders of fishes found in Nepal

Osteoglossiformes

Anguilliformes

Clupeiformes

Cypriniformes

Siluriformes

Cyprinodontiformes

Beloniformes

Synbranchiformes

Perciformes

Tetraodontiformes

Evolution of fishes

The evolution of fish began about 530 million years ago during the Cambrian explosion. It was
the time that the early chordates developed the skull and vertebral column, leading to the first
craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. During
the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish
known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant
lampreys may approximate ancient pre-jawed fish. The cyclomates which includes lamprey and
hagfishes may have split early on from other agnathans.

The earliest jawed vertebrates probably developed during the late Ordovician period. They are
first represented in the fossil record from the Silurian by two groups of fish: the armoured fish
known as placoderms, which evolved from the ostracoderms; and the Acanthodii (or spiny
sharks). The jawed fish that are still extant in modern days also appeared during the late
Silurian: the Chondrichthyes (or cartilaginous fish) and the Osteichthyes (or bony fish). The bony
fish evolved into two separate groups: the Actinopterygii (or ray-finned fish) and Sarcopterygii
(which includes the lobe-finned fish). During the Devonian period a great increase in fish variety
occurred, especially among the ostracoderms and placoderms, and also among the lobe-finned
fish and early sharks. This has led to the Devonian being known as the age of fishes.

Vertebrates, among them the first fishes, originated about 530 million years ago during the
Cambrian explosion, which saw the rise in organism diversity. The first ancestors of fish, or
animals that were probably closely related to fish were Pikaia, Haikouichthys and
Myllokunmingia. These three genera all appeared around 530 Ma. Pikaia had a primitive
notochord, a structure that could have developed into a vertebral column later. Unlike the
other fauna that dominated the Cambrian, these groups had the basic vertebrate body plan: a
notochord, rudimentary vertebrae, and a well-defined head and tail. All of these early
vertebrates lacked jaws in the common sense and relied on filter feeding close to the seabed.
These were followed by indisputable fossil vertebrates in the form of heavily armoured fishes
discovered in rocks from the Ordovician Period 500–430 Ma.

The first jawed vertebrates appeared in the late Ordovician and became common in the
Devonian, often known as the "Age of Fishes". The two groups of bony fishes, the actinopterygii
and sarcopterygii, evolved and became flourished. The Devonian also saw the demise of
virtually all jawless fishes, save for lampreys and hagfish, as well as the Placodermi, a group of
armoured fish that dominated much of the late Silurian. The Devonian also saw the rise of the
first labyrinthodonts, which was a transitional between fishes and amphibians.

The colonisation of new niches resulted in diversification of body plans and sometimes an
increase in size. The Devonian Period (395 to 345 Ma) brought in such giants as the placoderm
Dunkleosteus, which could grow up to seven meters long, and early air-breathing fish that could
remain on land for extended periods. Among this latter group were ancestral amphibians.

The reptiles appeared from labyrinthodonts in the subsequent Carboniferous period. The
anapsid and synapsid amniotas were common during the late Paleozoic, while the diapsids
became dominant during the Mesozoic. In the sea, the bony fishes became dominant.

General groups of fishes

Game/sport fishes of Nepal
The game fishes are those fishes which are generally caught by hooks in large rivers of Nepal. It
is for the purpose of recreational purpose. It is also called sport fishing. Approximately 118
varieties of fresh water fish live in the Himalayan waters of Nepal. Mahaseer is the most
commonly sought sport fish, besides mountain stream trout-like varieties and lowland water
species. Anglers can try their luck in Pokhara’s crystal clear lakes, or in west Nepal’s Bardia
Wildlife reserves where local agents can organizes guided fishing trips. While trekking or rafting,
bring along your pole, or give the locals’ method a try vita bamboo pole or crude fish trap.
Some important game fishes of Koshi and other rivers of Nepal:

Common/ Local Weigh

No Scientific Name Size Habitat Bait Season
Name t

Mahasheer/ 2.5 Large and deep

01 Tor putitora 50 kg Fish Mar-May./Sept-Dec
Sahar m rivers

DeepBodied Large and deep

02 Tor tor 3 m 50 kg Fish Mar-May/Sept-Dec
Mahasheer rivers

Copper Acrossocheilus Large and deep

03 6 m 5 kg Fish meat Mar-June/Sep.Nov.
Mahasheer /Katle hexagonolepis river

Fresh Water Large and Deep

04 Bagarius bagarius 3 m 100kg Fish Mar-May/Sept-Dec
Shark/Gounch rivers

Large and deep Worms

05 Jalkapoor Clupisoma gaura 5 m 1 kg Mar-May/Sept-Dec
rivers meat

Fresh Water Frogs

06 Anguilla bengalensis 3 m 18 kg River and pools Mar-May/ Sept-Dec
Eel/Raj Bam worms

River Catfish/ Worms Mar - May/ Sept-

07 Silondia silondia 5 m 2.5 kg River confluence
Bachawa meat Dec

Giant Murrel/ 2.5 River, stream and Worms, fish

08 Channa maurilus 15 kg Fe-May/Sept to Dec
Saur m lakes insects
 Amphipnous
River, stream,
09 Mud Eel/Hile Bam (Monopterus) 8 m 1.5 kg Meat Feb-Apr.
lakes and swamps
cuchia

Worm,
Feather Back/ Notopterus River, streams and
10 4 m 2 kg insects Feb-May / Sept-Dec
Patali notopterus lakes
meat

Worm,
River, streams and
11 Knife Fish/ Chitala Notopterus chitala 1 m 8 kg insects, Feb-May/Sept-Dec
lakes
meat

Fish,
1.8 River, streams and
12 Catfish/Buari Wallago attu 12 kg insects, Feb-June/Sept-Nov.
m lakes
meat

Tenger Catfish/ Fish, insects

13 Mystus seenghala 4 m 14 kg River streams Jan-June/Sept-Nov
Tengri meat

Important fish species as baits Predatory species

Common Name Common Name Scientific Name

Mahasheer and
01 Flying Barb Esomus danricus
Catfish

02 Sucker Head Garra gotyla Mahasheer

 Heteropneustes Catfish, Fresh Water
03 Singhe Catfish
fossilis Shark 'Buster'

04 River Barb Puntius chillinoides Copper Mahasheer

05 Gravel Catfish Glyptothorax telchitta Mahasheer

06 Torrent Minnow Barilius sps Mahasheer

Exotic fishes of Nepal

In biology, an exotic species refers to an animal species that is non-native. It is
introduced into an area where it does not occur naturally. The introduction of the
species may be deliberate or accidental. In Nepalese freshwater bodies, several fish species
have been introduced since 1956. The first exotic fish to be introduced in the country was the
Common carp (Cyprinus carpio) (Gubhaju, 2008). Rajbanshi (1982) reported seven exotic fish
species introduced for commercial uses; Shrestha (1995) reported ten exotic fish species;
GoN/MoFSC (2014) reported 12 exotic fish species whereas GoN/NPC (2019) reported eight
exotic species. Now, there are 15 exotic fish species in Nepal. Indigenous and exotic fish species
are farmed in Nepal. Terai is the site for carp, tilapia and catfish. Among carp the three major
Indian carps commonly farmed are Rohu (Labeo rohita), Catla (Catla catla) and Mrigal (Cirrhinus
mrigala). In addition, exotic carps namely Common carp (Cyprinus carpio), and Chinese carps:
Grass carp (Ctenopharyngodon idellus), Silver carp (Hypophthalmichthys molitrix) and Bighead
carp (Aristichthys nobilis) have been cultured since 1955/56. Among catfish mostly Clarias
batrachus, Clarias gariepinus and Pangasius pangasius are cultured. Recently goldfish (Carassius
auratus) was introduced as a recreational species in Nepal. In lower mid-hill or in cold water
zone, carp along with cool-water fish species such as Mahseer (Tor spp) and Katle
(Acrossocheilus hexagonolepis), Asla or cold water cyprinids (Schizothorax spp) and rainbow
trout (Oncorhynchus mykiss) can be cultivated. In Nepal rainbow trout (Oncorhynchus mykiss)
was introduced from India in 1968 and 1971 and from Japan in 1988. Recently, Nile tilapia
(Oreochromis niloticus), Java barb (Barbonymus gonionotus) have been introduced to study the
potential of their commercial production.

List of exotic fishes

S.N. Scientific name Common name
1 Barbonymus gonionotus Silver barb
2 Carassius carassius Goldfish
3 Catla catla Bhakur
4 Cirrhinus mrigala Naini
5 Clarias gariepinus African maghur
6 Ctenopharyngodon idella Grass carp
7 Cyprinus carpio Common carp
8 Hypothalamichthys molitrix Silver carp
9 Hypothalamichthys nobilis Bighead carp
10 Labeo rohita Rohu
11 Oncorhynchus mykiss Rainbow trout
12 Oreochromis mossambicus Tilapia
13 Oreochromis niloticus Tilapia
14 Pangasianodon hypophthalmus Pangasia
15 Salmo trutta Brown trout

Migration in fishes
Migration of fish is defined as a class of movement which involves a long journey to a definite
area for some purpose and impels the migrants to return to the region from which they have
migrated. The purpose of the journey is breeding and feeding.

Migration is a two-way journey. It includes emigration (outward journey) and immigration

(return journey or inward journey). The fishes are notable for migration for the purpose of
spawning. The inherent purpose of migration is not known.

Types of Migration: Myers (1949) has classified the following types of fish migration.

a. Diadromous migration:

When the migrations occur in between freshwater and marine environments.

Diadromous type of migration can be divided into following three-types.

(i) Anadromous migration: When migration occurs from sea to freshwater for
spawning, called anadromous migration; e.g., Atlantic salmon (Salmo salar), Hilsa
shad (Tenualosa ilisha), Toli shad (Tenualosa toli), Paradise fish (Polynemus
paradiseus), Flat head sillago (Sillaginopsis panijus), Sturgeon (Acipenser) and
Salmon (Oncorhynchus).
(ii) Catadromous migration: The journey of freshwater fishes to the sea for spawning,
called catadromous migration; e.g., Indian longfin eel (Anguilla bengalensis
 bengalensis), Shortfin eel (Anguilla bicolor bicolor), Common freshwater or European
eel (Anguilla anguilla, A. vulgaris), American eel (Anguilla rostrata).
(iii) Amphidromous migration: Migration of fishes from freshwater to sea and vice versa
and is not for the purpose of breeding but for the other purposes (e.g., food). The
amphidromous fishes migrate regularly at some particular stage of the life cycle.

b. Potamodromous migration: Migrations of fish that occur entirely in freshwater, called

potamodromous e.g., carps and trout. Trouts and carps travel long distances in large shoals in
search of suitable spawning grounds and return to feeding areas after spawning.

c. Oceanodromous migration: Migration which occurs entirely in sea, called oceanodromous

migration. Horizontal and vertical distribution is considered in oceanodromous migration. Many
fishes undertake short distance migrations throughout their life and some fishes like herrings,
cod, tuna and plaice, cover long distance migrations.

Causes for Fish Migration: Fish migration is related to several factors such as physical, chemical
or biological.

i. Physical factors: The physical factors include temperature of water, rainfall, quality of water,
water depths, pressure, light intensity, photoperiod, turbidity, tides and currents.

ii. Chemical factors: The chemical factors include pH of water, salinity, dissolve of O2 and CO2,
types of dissolved organic and inorganic substances, and taste of water.

iii. Biological factors: The biological factors are food, attainment of sexual maturity, endocrine
behaviour and competitors and predators.

Significance of migration

1. to find suitable feeding and spawning ground

2. for protection from predators
3. survive from extreme climatic conditions
4. increases genetic diversity
5. it is an adaptational characters for survival and existences

Accessary respiratory organ of fish

Accessory respiratory organs in fishes

Generally adult fishes depended chiefly on pharyngeal gills for aquatic respiration. However,
other devices also occur to supplement or replace gill respiration. All such additional respiratory
organs, other than gills, are known as accessory respiratory organs. The development of
accessory respiratory organs is found mostly in freshwater fishes of tropical region and very
rarely in marine fishes.

Sometimes the fishes of the tropical freshwaters and hill-streams develop accessory respiratory
organs to meet extra demand for oxygen, because depletion of oxygen occurs during summers
as the water level falls to a considerable degree. Accessory respiratory organs enable the fishes
to live in oxygen-deficient water, to aestivate over prolonged droughts in dry summer, to take
excursions on land or simply to meet extra demand for oxygen.

To overcome these adverse situations, accessory respiratory organs functionable in aquatic

and/or aerial environment have been developed in fishes. So the development of such
structures is essentially adaptive in native. Some accessory organs sub serve aquatic respiration,
while others aerial respiration.

Several types of accessory respiratory organs have been evolved in different

species of fishes. These accessory respiratory organs of fishes are as follows:

1. Skin or Integument:

In the eel, Anguilla anguilla, Amphipnous cuchia and in Periophthalmus and Boleophthalmus,
the skin is highly vascular and serves for exchange of gases as in frog, when the fish is out of
water. These fishes habitually leave the water and migrate from one place to another through
damp vegetation. During this period, the moist skin serves as an important organ in respiration.
They can respire cutaneously both in air and in water.
Since Amphipnous and Mastacembelus live in oxygen deficient stagnant water, the skin is of
little use for respiration but it plays an important role in extracting oxygen from air, when the
fishes are exposed in drying up muddy ponds, or when fish is moving out of water. The
glandular secretions of the skin protect it from desiccation in the air.

Median fin folds of many fishes are supplied with numerous blood vessels that help in
cutaneous respiration. Besides, the highly vascular opercular folds of Sturgeons and many cat
fishes serve as accessory respiratory structures.

2. Bucco-Pharyngeal Epithelium:

In most of the fishes, the epithelial lining of buccal cavity and pharynx is usually highly vascular
and permeable to gases in water. It may remain simple or may develop folds, pleats or tongues
projecting into the buccal cavity and pharynx to make it an efficient respiratory organ.

But in mudskippers (Periophthalmus and Boleophthalmus) the highly vascularised

buccopharyngeal epithelium helps in absorbing oxygen directly from the atmosphere. These
tropical fishes leave water and spend most of the time skipping or walking about through
dampy areas particularly round the roots of the mangroove trees. The old idea that the
mudskippers use the vascular tail as the respiratory organ is not supported by recent
ichthyologists.

3. Gut Epithelium:

In several fishes epithelial lining of certain parts of alimentary canal becomes vascular and
modified to serve as a respiratory organ. It may be just behind stomach (Misgurus fossilis) or
intestine (Lepidocephalus guntea, Gobitus (giant loach of Europe) or rectum (Callichthyes,
Hypostomus and Doras).

Fresh air is drawn through mouth or anus and after gaseous exchange the gas is voided through
the anus. In these fishes the wall of the gut is modified to perform the respiratory function. The
walls of the gut in these areas become thin due to the reduction of muscular layers.

4. Outgrowths of Pelvic Fins:

In American lung fish, Lepidosiren, during breeding time, the pelvic fins of male become
enlarged and grow filamentous vascular outgrowths which provide fresh oxygen to the guarded
eggs.

5. Pharyngeal Diverticula:
Pharyngeal diverticula are a pair of simple sac-like outgrowths of pharynx, lined by thickened
vascular epithelium and extending above the gills. In Channa (= Ophiocephalus), the accessory
respiratory organs are relatively simpler and consist of a pair of air-chambers.

These are developed from the pharynx and not from the branchial chambers as seen in others.
The air-chambers are lined by thickened epithelium which is highly vascularised. The air-
chambers are simple sac-like structures and do not contain any structure. These chambers
function as the lung-like reservoirs. In Channa striatus, the vascular epithelium lining the
chambers becomes folded to form some alveoli. The gill-filaments are greatly reduced in size.

In cuchia eel, Amphipnous cuchia, the accessory respiratory organs consist of a pair of vascular
sac-like diverticula from the pharynx above the gills (Fig. 17.8). These diverticula open anteriorly
into first gill-slit.

These diverticula function physiologically as the lungs. The gills are greatly reduced and a few
rudimentary gill-filaments are present on the second of the three remaining gill-arches. The
third gill-arch is found to bear fleshy vascular (respiratory) epithelium.

In Periophthalmus also, a small, shallow pharyngeal diverticulum lined with respiratory

epithelium (vascular epithelium) is present on each side of the roof of the pharynx.

6. Opercular Chamber Modified for Aerial Respiration:

In some species, the inhaled air is passed through the gill-slits into the opercular chamber
where it is stored for some time. The opercular chamber becomes bulged out in the form of
two little balloons in the hinder region of the head and after sometimes its walls collapse and
the air is passed out through the small external branchial opening. The membrane lining the
opercular chamber becomes thin and highly vascular to allow exchange of gases. This is seen in
Periophthalmus and Boleophthalmus.

7. Branchial Diverticula:

The outgrowths from gill-chambers form more complicated aerial accessory respiratory organs
than the simpler pharyngeal outgrowths in other fishes. Such air breathing organs are present
in Heteropneustes, Clarias, Anabas, Trichogaster, Macropodus, Betta, etc.

Important modifications in some of these species are described below:

(a) Heteropneustes Fossilis (= Saccobranchus):

This Indian catfish has a pair of long, tubular and dorsally situated air-sacs, arising posteriorly
from gill-chambers and extending almost up to the tail. They are highly vascular. The air is
drawn in and expelled out through pharynx.
(b) Anabas Testudineus:

The Indian climbing perch has two, spacious, suprabranchial cavities as dorsal outgrowths of
the two gill-chambers. Each cavity contains a special labyrinthine organ formed of much folded,
concentric bony plates developed from the first epibranchial bone and covered with thin
vascular mucous membrane. Margins of these plates are wavy and the plates are covered with
vascular gill-like epithelium.

Each branchial outgrowth communicates freely not only with the opercular cavity but also with
buccopharyngeal cavity. Air is drawn through mouth into suprabranchial cavities and expelled
through opercular opening. The fish is so dependent on atmospheric oxygen that it will drown if
denied access to surface to gulp air.

(c) Trichogaster Fasciatus:

The accessory respiratory organs in this species consist of a suprabranchial chamber, a

labyrinthine organ and the respiratory membrane. The suprabranchial chamber is situated
above the gills on either side as in Anabas, communicates with the pharynx by means of
inhalent aperture and with the exterior through the opercular chamber by means of an
exhalent aperture.

The labyrinthine organ develops from the epibranchial of the first gill-arch and is simpler in
structure than that of Anabas. It is in the form of a spiral organ possessing two leaf-like
expansions and is composed of loose connective tissue covered by a vascular epithelium.

The respiratory membrane lining the air-chamber and covering the labyrinthine organs consist
of vascular and non-vascular areas, of which the former possesses a large number of ‘islets’
containing parallel blood capillaries. The islets are believed to be derived from the secondary
lamellae of a typical gill-filament.

(d) Clarias batrachus:

The Indian cat fish, Clarias batrachus has the most complicated accessory respiratory organs.
The accessory respiratory organs are

(i) The suprabranchial cavity or chamber,

(ii) The two beautiful ‘rosettes’ or air-trees or arborescent organs or dendritic organs,

(iii) The ‘fans’ and

(iv) The respiratory membrane.

The suprabranchial chamber lies above the gills and is divided into two cup-like compartments
and is lined by a highly vascular respiratory membrane.

Two beautiful ‘rosettes’ or dendritic organs are present on each side and are supported by
epibranchials of the second and the fourth branchial arches. The first of these is smaller in size
and lies in the anterior compartment. Each is a highly branched tree-like structure supported by
cartilaginous internal skeleton. The terminal knobs or bulbs of each dendritic organ consist of a
core of cartilage covered by vascular epithelium showing eight folds in it.

According to Datta Munshi, each knob represents eight abbreviated and fused gill-filaments.
Some of the primary gill-lamellae of each gill-arch are fused so as to form a ‘fan’ or gill-plate.
Hence, there are ‘four ‘fans’ on either side and each consists of vascular and non- vascular
areas. The respiratory membrane lining the suprabranchial chamber is also composed of
vascular and non-vascular areas, of which the former show a large number of ‘islets’.

According to Datta Munshi (1961), the respiratory membrane has developed by the
abbreviation and fusion of the primary gill-lamellae and shortening of secondary gill-lamellae.
Well-developed exhalent and inhalent apertures are present for the suprabranchial chamber.
The fish rises to the surface of water and gulps in air, which from the pharynx enters into the
suprabranchial chamber through the inhalent aperture.

When the air enters the opercular cavity, it is directed into the suprabranchial chamber by the
action of the two fans on the second and third gill-arches. The exhalation of the air is effected
by the contraction of suprabranchial chamber and the movement of the fans. This creates a
partial vacuum in the suprabranchial chamber. The mouth is opened and the buccopharyngeal
cavity is enlarged to inhale air.

8. Air-Bladders:

Swim-bladder of higher bony fishes (teleosts) is essentially a hydrostatic organ. But in lower
bony fishes (dipnoans and ganoids), the air-bladder acts like a lung to breathe air and is truly an
accessory respiratory organ. The wall of bladder is vascular and sacculated with alveoli. In Amia
and Lepidosteus, the wall of the swim-bladder is sacculated and resembles lung.

In Polypterus, the swim-bladder is more lung-like and gets a pair of pulmonary arteries arising
from the last pair of epibranchial arteries. The swim-bladder in dipnoans resembles strikingly
the tetrapod lung in structure as well as in function. In Neoceratodus, it is single but in
Protopterus and Lepidosiren it is bilobed.
The inner surface of the ‘lung’ is increased by spongy alveolar structures. In these fishes, the
lung is mainly respiratory in function during aestivation because the gills become useless during
this period.

Like that of Polypterus, the ‘lung’ in dipnoans gets the pulmonary arteries from the last
epibranchial arteries. The swim-bladder of feather tail, Notopterus notopterus has a wide
pneumatic duct and a network of blood capillaries covered by a thin epithelium in its wall. This
helps in exchange of gases.

Functions of Accessory Respiratory Organs:

The accessory respiratory organs contain a higher percentage of oxygen. The fishes possessing
such respiratory organs are capable of living in water where oxygen concentration is very low.
Under this condition these fishes come to the surface of water to gulp in air for transmission to
the accessory respiratory organs. If these fishes are prevented from coming to the surface, they
will die due to asphyxiation for want of oxygen. So the acquisition of accessory respiratory
organs in fishes is an adaptive feature.

Further it has been observed that the rate of absorption of oxygen in such organs is much
higher than the rate of elimination of carbon dioxide. Hence, it is natural that the gills excrete
most of the carbon dioxide. Absorption of oxygen appears to be the primary function of the
accessory respiratory organs.

Origin and Significance of Accessory Respiratory Organs:

During development, the fifth gill-arch does not develop gill-lamellae, and its embryonic gill
material forms rudiments of the gill-arch, and aggregates to form a structure called the ‘gill-
mass’. The air-breathing organs or accessory respiratory organs develop from gill-mass. In some
species, the gill- arches other than the fifth arch, also take part in the formation of accessory
respiratory organs. The gill-lamellae which normally develop on gill-arches for aquatic
respiration, become modified to form the respiratory epithelium of the suprabranchial
chamber, dendritic organs and the air- sacs for aerial respiration. According to Singh (1993), the
air-sacs have evolved from the same basic material which has given rise to the gill in teleostean
fishes.

The accessory respiratory organs of Heteropneustes and Clarias, are modifications of the gills.
In these species, swim-bladder is either absent or greatly reduced. During the Tertiary and
Quaternary period of the Cenozoic era, the oxygen level of the atmosphere and of water was
reduced. Due to depletion of oxygen in rivers and swamps, the gills were unable to cope with
the requirements of the body. Hence, several teleostean species developed accessory
respiratory organs to absorb oxygen from air.
Most of the fishes possessing air-breathing organs or accessory respiratory organs are capable
of living in highly deoxygenated water of the swamps and muddy ponds infested with weeds.
They have been observed to gulp in air from the surface and to pass it to the accessory
respiratory organs. If prevented from reaching the surface, these fishes die due to asphyxiation.

This shows that the accessory respiratory organs are capable of maintaining life of the fish in
oxygen deficient water. It has been shown that the absorption of oxygen in the accessory
respiratory organs is much greater than the excretion of carbon dioxide. Hence, most of the
carbon dioxide is excreted in the gills and the chief function of the accessory respiratory organs
is the absorption of oxygen required for sustenance of life.

Scales in fishes

Fish scales are part of the Fish’s integumentary system, and these are originated from the
Mesoderm layer of the dermis.

A fish scale can be defined as the rigid plate that grows out of the skin of a fish. In most of the
fishes, the skin is covered with these protective scales. These scales also provide effective
camouflage through the use of reflection and colouration, as hydrodynamic advantages. The
scales vary in shape, size, structure, and extent that ranging from strong and rigid armour plates
in fishes like shrimp fishes and boxfishes, in some, it is microscopic or absent such as Eels, and
Anglerfishes.

Various types of Scales in Fishes:

Cosmoid Scales: Cosmoid scales are found only on ancient lobe-finned fishes including some
earliest lungfishes (Dipnoi) and Crossopterygii. They were probably derived from a fusion of
placoid scales. Cosmoid scales increase in size through the growth of the lamellar bone layer.

Placoid Scales: These are also called denticles. The placoid scales are structurally homologous
to the vertebrate teeth “denticle” with a central pulp cavity supplied with blood vessels, that is
surrounded by a conical layer of dentine, all of which sits on top of a rectangular basal plate
that rests on the dermis. Placoid scales cannot grow in size, but rather more scales are added as
the fish increases in size. The outermost layer is composed of vitrodentine, a largely inorganic
enamel-like substance. In Shark, the skin is almost covered by small placoid scales, supported
by spines. During forward movement in shark, these scales create tiny vortices which reduce
hydrodynamic drag and reduce turbulence, that makes swimming more efficient and quitter
compare to bony fishes. Placoid scales also serve a role in anti-fouling by exhibiting the Lotus
effect. These scales are found in cartilaginous fishes as such Sharks, rays, and chimeras.
Ganoid Scales: Ganoid scales are derived from cosmoid scales. Ganoid scales are covered with a
hard enamel layer like dentine and a layer of inorganic bone salt called ganoine instead of
vitrodentine. Ganoine is a characteristic component of ganoid scales. In appearance, it is glassy,
often multi-layered mineralized tissue, which covers the scales as well as the cranial bones and
fin rays in some non-teleost ray-finned fishes. For e.g. Sturgeons, paddlefishes, gars, bowfin,
and bichirs.

Cycloid Scale: It is a circular scale with a smooth texture and uniform with a smooth outer edge
or margin. They are most common in fishes with soft fin rays, such as Salmon and carps.

Ctenoid Scales: Ctenoid or toothed scales are like cycloid scales, but there is an exception that
they have small teeth or spinules (small spines or thorns) called ctenii found on their outer or
posterior edges. Because of the presence of these small teeth or spinules, ctenoid scales have a
rough texture.These fins are usually found in fishes that possess spiny fin rays such as Perch like
fishes.

Fins of fishes

Fins are one of the most distinguishing features of a fish and they have several different forms.
Two types of fins are found in most of the fish: median and paired fins. Median fins are single in
number which runs down the mid-line of the body. In fishes, median fins are dorsal, caudal and
anal fins while paired fins are pectoral and pelvic which are arranged in pairs homologous to
human arms and legs. Fins help to swim and maintain the balance of the body. Fins also help to
identify the fish species. Different types of median and paired fins are described below:

Dorsal Fin

This type of fin is located on the top or back of the fish which help the fish in quick turns or
stops. It also helps the fish against rolling. In fish, there are three distinct dorsal fins such as
proximal, central or middle, and distal dorsal fins. Some fish have two dorsal fins where the
central and distal fins are combined together.

Pelvic Fin

In fishes, a pair of pelvic fins are present which are located ventrally below and behind the
pectoral fins. In some fishes, they are situated in front of the pectoral fins (Cod family). This
type of fin helps in stability and slowing down the fish. Generally, fish use pelvic fins for moving
upwards and downwards in the water.

Anal Fin

The Anal fin is also known as cloacal fin which is located on the ventral side just behind the
anus. It supports the dorsal fin and stabilizes the fish during swimming and contrinols the rolling
motion.

Pectoral Fin

Pectoral fins are located on both sides usually just behind the operculum. It is homologous to
the tetrapod`s forelimbs. It provides supports during swimming. It creates dynamic lifting force
and also helps the fish to turn left or right.

Adipose Fin

They are soft fins and located between the dorsal and caudal fins, usually very near to the
caudal fin. It is mainly found in Salmonidae, Characins, and catfishes. This type of fin helps to
navigate the fish in rough water.

Caudal Fin

The caudal fin is the primary appendage which is used for locomotion in many fishes. The
caudal fin is also known as tail fin or a median fin which is usually homocercal or heterocercal.
Generally, it is a vertically expanded structure which is located at the caudal end of the body.
The base of the caudal fin is known as caudal peduncle with strong swimming muscles. In
general, caudal fin acts like a propeller while the caudal peduncle functions as a motor.

The caudal fin has two lobes such as dorsal epichordal and ventral hypochordal lobe which are
supported by the modified last three caudal vertebrae. The shape of the caudal fin may vary in
different species from rounded to pointed, notched, emarginated, truncated, etc. It is used to
identify the fish species. Generally, fish use it for forwarding propulsion and speed.

The caudal fin of the adult fishes may be grouped into three categories:
Protocercal Caudal Fin

It is the most primitive type of caudal fin where the straight vertebral column divides the caudal
fin into two equal lobes such as upper lobe and lower lobe. In this case, the upper lobe is
known as epichordal or epicaudal and the lower lobe is called hypochordal or hypocaudal lobe.
A series of rods are arranged around the central axis of the caudal region, which support the fin
membrane. Undoubtedly, during the developmental period, the caudal fin of all fishes passes
through the protocercal stage. This type of fin is found in cyclostomes and the living
dipnoans(lungfishes).

Heterocercal Caudal Fin

The heterocercal tail is sometimes called the shark-tail type of caudal fin. Elasmobranch
(cartilaginous fish) and some primitive type of bony fishes contain this type of fin. This fin has
two unequal lobes where the upper smaller lobe is known as epichordal lobe and a much larger
lower lobe is known as hypochordal lobe. In this case, the hind end of the vertebral column
becomes bent upwards and continues almost up to the tip of the fin.

Homocercal Caudal Fin

Most of the higher teleosts have homocercal caudal fin. It has superficially symmetrical and two
equal sized lobes such as upper epichordal and the lower hypochordal lobe. Internally, this tail
is asymmetrical and the hinder end part of the vertebral column is greatly shortened and
turned upward. In this case, the vertebral column does not touch the posterior limit of the fin.
Varieties of Caudal Fins

The internal and external structure of caudal fin varies which depends on the swimming habits
of the fish. Generally, these variations involve special modification of the vertebral column.
Following seven main types of caudal fins are found in fishes:

Lunate or Crescentic: It is used for Continuous long distance swimming. e.g. Tuna

Forked: It is used for rapid swimming, e.g. Herring, Mackeral.

Emarginate: e.g. Trout, Carp, Perch.

Truncate: It aids in turning quickly. e.g. Flounder

Rounded: It is used for slow swimming, accelerating, and maneuvering. e.g. Turbot and Lemon-
Sole.

Pointed: e.g. Gobby

Double emarginated
 Parental care in fish

Deposition of eggs in suitable places:

In some fish species, the eggs are deposited at their own designed suitable places. They choose
places like:

Deposition of eggs in sticking covering: Carps laid with special sticky coverings for the
attachment to each other or to stones, weeds, etc. Yellow perch deposits eggs in a rope-like
structure, a long floating membrane works for eggs to hold together. Flying fishes, garfishes,
skippers, etc. secrete a sticky thread-like substance from the kidney by which eggs remain
attached.

Laying eggs at suitable places: The anadromous fishes like Acipenser, Onchorynchus, etc. lay
their eggs in suitable places for spawning. They lay their eggs in pits and cover them with gravel
and desert them.

Depositions of eggs on dead shells of bivalves: Fishes like the female of European bitterling
(Rhodeus amarus), females like cyprinid family deposits their eggs in dead shells, water
Mussels.

Scattering of eggs over aquatic plants. Examples of such parental care in fishes as such pikes,
carp, the eggs are scattered and attached to aquatic plants.

Self-made nest for deposition of eggs: Some species of fish prepare different types of nests for
the deposition and protection of their eggs. These nests may be Basin-like, Circular nest, Hole,
burrow nest, Barrel shaped to nest, Cup-shaped nest, Floating nest, and Foamy nest. Examples
of different types of nests made during parental care.

Female Darter makes basin-like depression and deposits eggs in it. The eggs are fertilized by the
male and covered by a sticky secretion from its kidney. These sticky eggs stay attached to stone
till hatching. The Cichlid fishes build a basin like a nest and it is guarded by both parents. Male
Bowfin (Amia calva) makes a circular nest. The male African lungfish (Protopterus) makes a nest
in the form of a deep hole in swampy places along the river bank. The male Mormyrids
(Gymnachus) make a floating nest in aquatic vegetation. The male Betta fish (Fighting fish)
makes a nest by blowing bubbles of air and sticking mucus that adheres and forms a floating
mass of foam on the water surface. The fertilized eggs are collected in the mouth of males. The
male stays and guards the eggs and fights till death to protect them.
Concealing eggs and young inside or on their body:

This is the highest degree of parental care, found in viviparous species. In some species to
provide maximum protection, internal incubation and they give birth to young ones.

Example of Viviparous parental care:

Viviparous Elasmobranchs: The viviparity is found in most shark families. Fishes such as
Scoliodon, Mustelus, etc, eggs develop in the uterus, the mucus lining of the uterus forms fluid-
filled protective compartments, for each of the embryos. Each embryo gets nourishment in the
form of embruotrophe or uterine milk, from uterine tissue through the yolk sac placenta.

Viviparous bony fishes: Teleost species like gambusia, poicilia, etc also show internal
fertilization and the young one develops in the ovary but is free from its wall. Shiner perch
fertilizes eggs in the ovarian follicle and is very soon released into the ovary cavity. These are
nourished by a secretion from the ovary. The young are retained in the ovary till they become
sexually mature.
Care of independently swimming young one’s:

In some species, parental care does not stop with caring for eggs. In these families the fishes
care for young ones, placing them in a safe place and protect from predators and enemies. For
example, the families such as Gasterosteridae, Centachidae, and Ictaluridae. Some fish species
keep eggs and young in their body or mouth or brood pouches.

Example of Concealing eggs inside the body:in cichlids, the female keeps the fertilized eggs in
their mouth.In Catfishes (Arius) and cardinal fishes, the male carries eggs and young ones in
their mouth. During this period the male fish does not take food. The male Kurtus (nursery fish)
of new guinea, carries eggs on the forehead, held in the cephalic hook.

Some parents keep eggs in Brood pouches: like sea horses (Hippocampus). In sea horse, the
fertilized eggs are transferred by the female into the brood pouch of the male. The brood pouch
is found on the lower abdomen. For several weeks the male hippocampus provides nutrients
and oxygen to the fertilized eggs.

Male pipefishes (Syngnathus) a brood pouch is formed between two flaps of skin, on the
underside of the body. The female lays eggs on these brood pouches.