Marine Vertebrates

LEARNING MODULE:
Chapter 5

Allyn Duvin S. Pantallano, Ph.D.

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Disclaimer
This module is for the class use or purposes only and should not be shared without
prior consent. Contents of the module were sourced out from various references such as
online presentations, articles and related books. The whole content is not claimed by the
author as solely from her own collective ideas but rather from the references resources.
Thus, this module does not intend to infringe on any copy right claims.
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UNIT II: Biology of Fishes

CHAPTER 1 Food and Feeding habits

LEARNING OUTCOMES

At the end of this chapter, the students should be able to:

1. Know and identify the different kinds of food fishes eat
2. Know their feeding habits

Food and feeding habits of fishes is an interesting part of the biology to understand
the type and nature of food they consume and their place in the ecosystem as well.
Feeding is essential for growth, maintenance and reproduction in fishes. The fishes
exhibit feeding habits from simple filter feeding to highly predatory life. The feedings
habits of fishes are assessed by:

1. Field observation
2. Analysis of gut contents
3. Laboratory experiments

Categories of food based on availability

Food available in the nature can be classified as plankton, nekton, benthos and detritus.

1. Plankton : Plankton is microscopic organisms with little or no power of locomotion.

They drift at the mercy of water currents. They are broadly grouped into 2 categories
depending on the presence or absence of plant pigments.

1. Phytoplankton – having chlorophyll in the cells e.g. Diatoms,

Dinoflagellates
2. Zooplankton – without chlorophyll e.g. Copepods, crustacean larvae, other
microscopic invertebrates

Nekton : Nekton are actively swimming organisms capable of independent movement.

They actively maintain their position despite the drifting water currents. They may be
present at the surface (pelagic) or at the bottom (demersal) of water column. e.g. fishes,
shrimps, cuttlefishes and squids.

Benthos : These are bottom dwellers with little (sessile) or no movement (sedentary).
They are mostly invertebrates living on the substratum. e.g. annelid worms, bivalves,
gastropods etc., benthos are of two types i) phytobenthos and ii) zoobenthos.
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Detritus : It is the dead and decayed organic matter of both plant and animal origin.
Decomposing bits of leaves, twigs, barks, water plants and animals form detritus. It can
be in particulate or suspended form associated with rich microbial flora and fecal matter.

Categories of food based on importance

1. Main or basic food : Food which is normally eaten by the fish or the most
preferred food is the main food.
2. Occasional or secondary food : When the main food is not available, fish
feed on other available food temporarily. This type of food is called occasional
or secondary food.
3. Incidental food: This type of food that occasionally enters the gut of a fish
along with main food items.
4. Emergency or obligatory food : This is the one which fish takes in the
absence of basic food to maintain physiological activities. It is taken when
there is no alternative food available.

Further categories

Fishes are categorized based on their dependence on food type, preference,

position in water column and feeding types.

Dependence on food type : In nature, the type of food available does not remain the
same throughout. Therefore, fish may change over to other available food or restrict to a
limited type and the fishes are classified as:

1. Euryphagic : Fishes feeding on a mixed diet with no preference to any

certain type.
2. Stenophagic : Fishes feeding on limited kinds of food.
3. Monophagic : Fishes consuming only one kind of food.

Dependence on food preference: Depending on the food they take fishes are classified
as the following:

1. Herbivorous :A number of fishes feed on unicellular algae, filamentous

algae, and also higher aquatic plants. If the plant material in the diet is 75%
or more the fishes are considered to be herbivorous. i.e. Labeo fimbriatus,
L. rohita, Ctenopharyngodon idella, Oreochromis mossambicus .
2. Detritivorous : Those feeding mainly on detritus i.e. Labeo calbasu, Mugil
cephalus.
3. Omnivorous : Those feeding on both plant and animal matter. The food of
these fishes consists of varying percentage of plant and animal matter and
they form a link between herbivorous and carnivorous fishes. i.e. Etroplus
suratensis, Cyprinus carpio, Tor putitora, Cirrhinus mrigala, Clarias
batrachus, Heteropneustes fossilis.
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4. Planktivorous : Feeding mainly on plankton both phyto and

zooplankton i.e. catla - zooplankton feeder , silver carp - phytoplankton
feeder.

5. Carnivorous : Feeding on prey organisms. The examples of carnivorous

fishes are: Wallago attu, Mystus singhala, Channa striatus.

The carnivorous fishes may be:

1. Insectivorous: Mainly feeding on insects. i.e. Trout.

2. Carcinovorous: Mainly feeding on crustaceans. i.e. Black bass
3. Malacovorous: Feeding mainly on molluscs like snails/clams i.e. Black
carp
4. Piscivorous: Feed on fish other than its own species. But, generally they
prey upon small fishes of other species rather than their
own i.e. Barracuda.
5. Larvivorous: Feeding mainly on insect and crustacean larvae/fish
larvae. i.e. Gambusia affinis.
6. Cannibalistic: Feeding on the young ones of the same
species i.e. Channa marulius, Lates calcarifer.

Dependence on their position in the water column: Depending on the position in the
water column they occupy, the fishes are categorized as:

1. Surface feeders : Fishes which live in the surface and feeding mostly on
the food available in the uppermost layer of water. e.g. catla feeding on
zooplankton and silver carp feeding on phytoplankton.
2. Mid-water or column feeder : Fishes which live in the mid water or column
water and feeding in the middle layer are called mid-water or column
feeders. e.g. rohu feeds on plant matter including decaying vegetation.
3. Bottom feeders : Fishes which live at the bottom and feed on the benthic
fauna. e.g. mrigal feeds on detritus and decaying vegetation and common
carp on molluscs and chironomids in the bottom mud.

Dependence on feeding types: In composite fish culture, these feeding habits of fishes
are advantageous to utilize the available food in water body.

1. Predator. Predator fish are the one which feed on macroscopic animals. These
fishes have well developed teeth for grasping and holding to seize their prey
organism firmly. The predatory fishes have large mouth, reduced gill rakers, fairly
large stomach and short intestine. The stomach is provided with acid secretions to
digest the proteinaceous food material. (e.g. ribbon fish, seer fish, barracuda etc.).
Normally predators feed during the day time and while requires acute vision. But
in case of deep sea predators, eyes are not well developed, so they locate their
prey by smell, taste, touch and by lateral line sensory system and chase with large
gaping mouth.
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2. Grazers. Grazing type of feeding is comparable to grazing behavior of cattle and

sheep. Grazing fishes take their food by bites on a large spread of food organisms.
The grazers normally go in groups i.e. Butterfly fishes (Chaetodontidae) and Parrot
fishes (Sparidae) which are seen in coral reefs, feeding on coral polyps and on
algae among the reef. Lepidophagus grazing is a special type of grazing where the
fishes pluck the scales from other fishes. i.e. Cichlids. In hatcheries some fishes
exhibit strange type of graze-feeding behaviour. i.e. Trout and Salmon tear off
body parts from other fishes especially fins. In such fishes, mouth is pointed and
provided with plucking incisors.

3. Strainers. They strain or sieve water to get the food material for which their gill
rackers are modified. The gill rackers are numerous, elongated and closely set in.
With the help of such gill rakes, they filter large volume of water to obtain small
sized planktonic organisms. In plankton feeders, the food is selected by size not
by kind or type i.e. Sardine, Silver carp and Mackerel feed on copepods, but they
may take other plankton of the size of the copepods.

This type of feeding habits is seen in the case of bottom dwelling fishes. Here the
fishes swallow their food by sucking the desired food or food containing material.
In such fishes, lips are modified for sucking purpose. They suck food by either
segregating the food before sucking in or ingest along with unwanted materials
into the mouth and segregate it inside the mouth e.g. sturgeon. On account of
sucking habit many demersal fishes accumulate unwanted food material in large
quantities in the stomach e.g. catfish

4. Parasitic. Parasitism is a specialized mode of feeding and fishes are no exception

to this highly evolved feeding habit. The parasitic fishes, feed on the body fluids of
the host fish. Here the mouth is well developed and adapted to hold on to the host
and pierce and suck the body fluid. e.g. lampreys and hag fishes have terminal
mouth which is surrounded by short barbles to pierce the host fishes. Interestingly,
in deep sea angler fish (Ceratias), males which are smaller in size are parasitic on
the large bodied female fish. Shortly after hatching, the male finds a female and
attaches by its mouth to female are body and the female responds by developing
a fleshy papilla, from which the male can absorb nutrients. Once attached to the
papilla, the male takes no free living food at all.

Feeding adaptations

The diversity in feeding habits of fishes is the result of structural adaptations. The
structural adaptations for specialized feeding are seen in relation to lips, mouth, teeth, gill
rackers, digestive tract and sensory organs.
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Lips

Among fishes, there are Jawed (Gnathostoma) and Jawless (Agnatha) fishes. In
jawless parasitic lampreys (Petromyzontidae) and hag fishes (Myxinidae), the suctorial
mouth serves as hold fast organs for attachment to the host and as food remover from
the host. In jawed fishes, the mouth is terminal for biting the food. The fishes which take
large piece of food at a time do not have modified lips (all carnivorous fishes. Suctorial
feeders (suckers) have an inferior mouth and fleshy lips. The lips of Sturgeons and
suckers are mobile and described as plicate (having folds) or papillose (having small tufts
of skin or papillae). Lips not only help in feeding but they act as hold fast organs in rapidly
flowing mountain stream e.g. loaches. In addition, the suctorial feeders also have barbels
around the mouth. The sensory organs present in these barbels help in locating the food.

Mouth

Among the grazers and suctorial feeders, there exist not only specially developed
lips but also adaptations on other mouth parts. The trumpet fishes (Aulostomidae), the
cornet fishes (Fistulariidae) and the pipe fishes (Syngnathidae) as well as many butterfly
fishes (Chaetodontidae) of coral reefs have mouth that resembles elongated beak. With
elongated tube-like mouth, feeding in the case of trumpet fishes may be by suction and
in cornet fishes and Pipe fishes, it may be selective grazing action with sharp teeth. In
butterfly fishes, the long snout enables to reach into small cervices of the corals for picking
the food.
Predator fishes such as the Dories (Zeidae), certain Wrasses (Labridae) and the
European bream (Cyprinidae) can form temporary tubes which help in swallowing their
prey from close range by forward extension of the jaws. A peculiar modification of jaws is
shown by the half beaks (Hemiramphidae), in which the lower jaw projects into a beak
while upper jaw is small, as a result mouth opening lies above. Owing to this modification,
the beak helps in feeding at the surface of water.

Teeth

These are the major structures showing outstanding modifications for feeding. In
bony fishes there are three sets of teeth in jaws, mouth and pharynx. Teeth in the jaws
are those on the maxillary and pre-maxillary (upper jaw) bones, and on the dentaries
below (lower jaw bone). On the roof of the oral cavity, teeth are borne by the vomer,
palatine and ecto-pterygoid bones on each side. On the floor of the mouth, often tongue
has teeth on it. Similarly, the teeth in the pharynx occur as pads on various gill arch
elements in many species (Cyprinidae, Catostomidae). Many predatory fishes have teeth
like modifications on the inner surface of the pharyngeal arch e.g. northern pike (Esox
lucius).

Based on their shape and location, teeth in jaw are canine (fang like), incisor
(frontline cutting), molariform (grinding with flattened surface), cardiform (short, fine and
pointed arising from a pad), and villiform (elongated teeth that resembles the intestinal
villi).
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There is a strong relation between dentition, feeding habit and the food eaten by
fishes. Predators such as barracuda, ribbon fishes, silver bar have sharply pointed teeth
which help in grasping, puncturing and holding the prey. In skates (Rajidae) and in drums
(Sciaenidae), there are grinding (Molariform) teeth in oral or pharyngeal cavities. They
feed on snails, clams and hard bodied crustaceans. Razor like cutting teeth (Incisors) are
seen in predacious fishes like piranha of the Amazon and barracuda of warm seas. Teeth
are generally absent in plankton feeders and in some of the more generalized omnivores.

Gill rackers

The gill rackers arise from gill arch which supports the gill filaments. They also
protect the tender gill filaments from abrasion by the ingested materials that are coarse
in texture. The gill rackers are specialized in relation to food and feeding habits. In
plankton feeders, the gill rackers are numerous, elongated and closely set in for straining
the water efficiently. In many predatory fishes, which feed on larger prey, the gill rackers
are reduced or absent. Fishes which have intermediate feeding habit, the gill rackers are
of moderate size.

Digestive track

It includes oesophagus, stomach and intestine which exhibit various modifications in

relation to feeding habits.

Oesophagus is a highly distensible muscular tube which can accommodate

anything that a fish has ingested.

Stomach. It shows various modifications especially with respect to shape. In

piscivorous fishes, the stomach is typically quite elongate e.g. gars and barracuda.
In omnivorous species, the stomach is sac like. In some fish, stomach is modified
into grinding organ and reduced in size, but the inner wall is greatly thickened and
muscularized like a gizzard e.g. mullets, sturgeons, gizzard shads. In fishes which
devour a huge meal, the stomach is highly distensible, as in Bombay duck. A
remarkable modification of stomach exists in the puffers (Tetraodontidae) and
porcupine fishes (Diodontidae) which inflate it with water or air to assume an
almost globular shape. True stomach is not seen in all the fishes. In roach, a plant
eating cyprinid, the oesophagus directly leads to intestine, so also in parrot fishes.
True stomach is also absent in plankton feeders. The presence or absence of
stomach is not related to feeding habit but to whether or not the fish have structures
for grinding or triturating the food.

Intestine. The absorption of food material takes place in intestine. The length of
intestine is shorter in carnivores and much elongated and arranged in many fold in
herbivores (very long and highly coiled in rohu and mrigal while the omnivores
show an intermediate condition.
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Stimuli for feeding

Two kinds of stimuli are involved in feeding:

1. Factors affecting the internal motivation or drive for feeding include season,
time of day, light intensity, time and nature of last feeding, temperature and
any other internal rhythms that may exist.
2. Food stimuli perceived by the senses like smell, taste, sight and the lateral
line system that release and control the momentary feeding act. The
interaction of these two groups of factors determines when, what and how
a fish will feed.

Detection of food

Fishes are able to detect food by both physical and chemical senses by sensory
organs which orient the fish towards food. Based on the sensory responses, fishes are
categorized as:

Night feeders : Detecting the food by smell and taste e.g. catfishes.

Sight feeder : The fishes are active during the day time and their vision and
position of eyes is important in the feeding habit. Sight feeders feed intensely
during day but it does not mean that they don’t feed at during the night.

Surface feeders like minnows, eyes are oriented upwards above the mid lateral
line. This gives better view of objects at the surface of water.

Bottom feeders like cat fishes, the eyes are present below the mid lateral line and
they can able to see the bottom area better. The fishes which feed during day as
well as at night have well developed lateral line that helps in detection of food.
Lateral line system is well developed in blind cave fishes and deep sea fishes.

Other senses such as olfaction and taste also play a role in locating the food
available at a distance. Many fishes which feed by sight have acute olfaction.
Gustatory sensation enables the fish in the final selection and swallowing the food.
The taste buds are concentrated in the mouth, barbels, snout and sometimes on
the lips.

Selection of food

Not all the food materials that enter the mouth are swallowed by the fish. The final
selection takes place in the mouth and pharynx especially in bottom dwelling fishes.
These fishes feed on extraneous matter and it is subjected to selection inside the mouth.
This selection is made by various sensory structures situated in mouth, tongue, gill
rackers, epibranchial organs, and tissue surrounding the pharynx. The unwanted material
is thrown out and in many instances unwanted food is also thrown out from the gill
openings. In some cases, they throw the unwanted food by coughing action. Normally gill
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rackers, pharyngeal teeth, bristles and epibranchial organ serve as mechanical structures
in retaining or rejecting the food.

How much food fish feed on?

Generally herbivores and plankton feeder are continuous feeders and they feed all
through the day unlike carnivores. Food of plant origin has poor nutritional protein content
but more water and high fiber. In order to balance the nutritional requirements, herbivores
have to feed on large quantity e.g. grass carp. Conversely, the carnivores feed on animal
matter which is rich in nutrients; hence they take a small meal. Even in some carnivores,
the nature of food varies; most of the bottom carnivores feed on crustaceans and mollusks
which have more indigestible matter in the form of shells. So their intake is larger as
compared to piscivorous fishes.

Feeding periodicity and variations

Depending on the quantity of food consumed, the frequency of feeding varies from
few hours to 4-6 days e.g. deep sea fishes. Small feeders have more feeding frequency.
In addition, the factors regulating the feeding periodicity are: seasons, migratory cycles,
reproductive activity, age and size of fish.

Seasons

Seasons influence feeding frequency especially in temperate waters. Feeding

periodicity is related to water temperature and metabolic rate. Annual cycle of
temperature variation is more pronounced in temperate waters, where feeding frequency
is more in summer.

Migratory cycle and reproductive activity

In temperate waters, some fishes exhibit feeding patterns in relation to migration.

The fishes which undertake migration for breeding eat intensively and accumulate reserve
food material which enables development of gonads e.g. salmon. The younger fishes are
more active so they feed more than the older ones.

Amount of food consumed daily

Amount of food consumed daily depends mainly on the quantity of food and size of
fish. The amount of food consumed is related to metabolic rate which is in turn closely
related to the temperature. In tropical waters, temperature being higher the metabolic rate
and food requirement is also higher. Therefore, fishes living in tropical water feed on large
quantity of food e.g. tuna. The fishes living in temperate waters feed on small quantity of
food as the temperature is lower and the metabolic activity is also low e.g. Salmon.
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Size of fish

Small fishes consume more food when compared to large sized fishes. This is
related to body weight. For example, small fishes which weigh about 2-5 grams consume
6-10% of their body weight per day, whereas bigger fishes which weigh about 30 gm or
more feed on 2-3% of their body weight per day. The rate of consumption is also related
to the condition of the fish. In diseased fishes the rate of consumption is very low. During
spawning season, most of the fishes stop feeding due to the enlargement of gonads.
Interestingly, european eel (Anguilla) stops feeding after maturation as a result, digestive
system degenerates and finally it dies after spawning.