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2438 FISH/Demersal Species of Tropical Climates

Demersal Species of Tropical tagging data. But nowadays scientists are employing
the application of modern electronic tagging tech-
Climates niques, whereby perfect positional information can
G D Khedkar and B V Jadhao, Dr Babasaheb be derived after retrieval of the tag. The traditional
Ambedkar Marathwada University, Aurangabad, tagging experiments have provided evidence of
Maharashtra, India migrations across deep-sea areas. Even transatlantic
N V Chavan, Adarsha College, Hingoli, Maharashtra, crossings have been recorded for cod. Although such
India basin-wide migrations may not occur regularly, they
C D Khedkar, College of Dairy Technology, Warud certainly show that demersal fishes are quite capable
(Pusad), Maharashtra, India
of considerable movements over long distances.
Copyright 2003, Elsevier Science Ltd. All Rights Reserved.

Sensory Systems in Demersal Fishes of


Introduction Tropical Climates
0001 Demersal fishes have been a major source of protein The shelf environment is generally well-lit and most 0003

for people all over the world for thousands of years. demersal fishes have well-developed vision. In add-
Worldwide, about 85% of the total continental ition demersal fish may use mechanoreceptors (sense
shelf area has sandy or muddy substrate. Only about of pressure, motion, and sound), chemosensory
6% is rocky or gravelly, and the remaining areas are (olfaction and gustation) and electrosensory (i.e., abil-
coral reefs or shell beds. However, there are both ity to detect electric fields) systems. Different systems
latitudinal and depth-related patterns. Corals are may play essential roles during different stages of
almost entirely confined to low latitudes where or- important processes such as prey detection, capture
ganically enriched muddy areas are also most exten- and handling, predator avoidance, or reproduction. In
sive, particularly near the mouths of major rivers general, mechanoreception by the acousticolateralis
or below highly productive upwelling areas. Sandy, system (lateral line organ and ear) tends to be most
rocky, and gravelly sediments are more common important for the detection of predators, conspecifics,
at high latitudes. The tropical waters contain a and other physical disturbances in the environment.
greater number and diversity of fish species than Olfaction is very important to benthofages that have
waters of the higher latitudes. Demersal fish of to search the substrate for food, whereas planktivores
economic importance are concentrated on the contin-
ental shelf and on the continental slope down to
about 500 m. They live on or near the bottom of the
ocean. A regular landing from a deep-sea trawling Table 1 Total world catch of major demersal fish species tbl0001

(1990–2000)
session would include catfish, croakers, soles,
groupers, shinynose, snappers, mullets, and mega- Scientific name Commonname Catch
lops. World catches of demersal species increased (tonnes)
rapidly during the first three-quarters of the 20th Ammodytes spp. Sand eels 1 003 343
century. Since then the catches from some stocks Ariidae spp. Sea catfish nei 242 815
have declined, due to overfishing, while other previ- Cantherhines spp. Filefish 234 446
ously underexploited stocks have increased their Elasmobranchii Sharks, skets, rays 337 819
yields (Table 1). The limits of biological production Gadus macrocephalus Pacific cod 425 467
Gadus morhua Atlantic cod 2 317 261
have probably been reached in many areas. In Macruronus magellanicus Patagonian grenadier 230 221
order to maintain the fisheries and to protect the Macruronus novaezelandiae Blue grenadier 255 421
ecosystems which support them, careful management Melanogrammus aeglefinus Haddock 273 459
is needed. Merluccius capensis Cape hakes 266 854
Merluccius gayi South Pacific hake 197 911
Merluccius hubbsi Argentine hake 526 573
Migration in Demersal Species of Tropical Micromesistius poutassou Blue whiting 628 918
Climates Nemipterus spp. Threadfin breams 186 201
Pleurogrammus azonus Atka mackerel 237 343
0002 Diurnal migrations, either vertically or horizontally, Pleuronectiformes Flatfishes nei 289 551
are common in demersal fishes. More fascinating, Pollachius virens Pollock 385 227
however, are the long-distance seasonal migrations Sciaenidae Croakers 492 528
associated with reproduction, feeding, and sometimes Sebastes spp. Atlantic redfishes 318 383
Theragra chalcogramma Alaska pollock 3 182 645
overwintering. Many examples were already well
Trisopterus esmarkii Norway pout 299 145
described several decades ago, based on traditional
FISH/Demersal Species of Tropical Climates 2439

and piscivores tend to rely more heavily on vision. Demersal sharks are either oviparous or vivipar- 0006

Gustation is most important after capturing prey. ous, whereas rays and chimaeroids are oviparous,
Benthofages may have elaborate adaptations of their attaching their few large encapsulated eggs to debris
olfactory system. In addition to a well-developed or macroalgae. All species produce rather few young
nose, they may have barbells on the snout (e.g., in each batch, i.e., from a couple to a few tens of eggs/
gadids, zoarcids) and modified and extended first juveniles, and lifetime production is very low com-
rays of their pectoral fins (e.g., Triglidae) densly pared with most teleosts. Upon hatching or birth, the
packed with olfactory sensory cells. The electrosen- young resembles the adults. Spawning seasons vary in
sory system is particularly well developed in some duration, and the general rule is that the more sea-
sharks and rays, and they may primarily use this sonal the production cycle, the more fixed and limited
system for prey location, but also for orientation and is the spawning season. High-latitude fishes and those
navigation. living in monsoon or upwelling regions have com-
paratively short spawning seasons, whereas tropical
and subtropical fishes have protracted seasons or
Life History Characteristics of the
year-round reproduction. Larvae hatched from small
Demersal Fishes of Tropical Climates
pelagic eggs have a yolk sac that may provide suffi-
0004 Life history strategies are best examined in the con- cient nutrition for a few days or weeks, after which
text of the assemblage or community within which they depend on exogenous feeding, usually on small
the fish operates. No species lives in isolation, but crustacea such as copopod nauplii. The mortality in
interacts with and influences the activities of all the this perod is very high indeed, and only a minute
other species it regularly encounters. Ecological fraction of the total number of eggs spawned will
understanding ultimately grows out of the study of eventually result in a surviving demersal juvenile.
communities, and this perspective is the one most The teleost postlarvae pass through metamorphosis
deep-ocean ecologists have adopted. Comparative upon reaching a certain size. Characters such as fins
study of several systems, when possible, can be very and juvenile pigmentaion develop, and sense organs
insightful. The samples required for this sort of study, also become fully developed. In flatfishes, the eye
however, are not easily come by. Large numbers are migration occurs. In general, the larva is transformed
required, which, in the best case, should encompass into a fish which is morphologically and behaviorally
most life stages and time series over seasons, over adapted for demersal life. Associated with this change
years, and over space. A single sampling gear rarely is the settling on the seabed, at least for parts of the
suffices, and even a suite of different nets, for diurnal cycle. Settling areas are often, but not always,
example, might also be complemented with cameras separated from the feeding areas of the older fish.
and other ‘snapshot’ devices. For deep-sea fishes, it is This reflects the different habitat and food require-
hard to find any area where such requirements are ments of juveniles and adults, but may also reduce
met. One that comes somewhat close is in the deep cannibalism. In the majority of teleost species, the
abyssal region of the eastern Atlantic. demersal nursery areas are shallower than feeding
0005 Most demersal teleosts (bony fishes) are oviparous grounds of older conspecifics. Estuaries and offshore
and produce free-floating eggs. Mating and spawn- shoals are typical nursery areas, also rocky shores
ing happen at the same time, often in mid-water. where the substrate and macroalgae offer protection
Some notable exceptions are the viviparous redfish and a variety of prey. Tidal flats and sandy beaches
(Sebastes) that release numerous pelagic larvae, and are typical habitats of juvenile flatfish. A gradual
the wolfish (Anarhichadidae) which deposits a cluster ontogenetic shift in depth distribution happens as
of large eggs that is guarded until hatching. Sand eels the juveniles grow larger.
(Ammodytidae) also have demersal eggs but do not The expected longevity of demersal species of trop- 0007

protect the eggs. Most teleosts have no parental care ical climates varies greatly. A general pattern is that
other than making sure that they mate and spawn longevity increases with increasing adult size, but
in an area and habitat that enhances the survival there are many exceptions to this rule. Some species,
probability of their eggs, larvae, and pelagic juveniles. i.e., redfishes, can probably live for at least 30–40
Most pelagic teleost eggs are small, i.e., a few years, but this is unusual for shelf species. Small
millimeters in diameter, and fecundity is high, i.e., species such as Norway pout may live for 5–6 years
thousands to millions of eggs per female. Batch at most, but in exploited areas this seldom happens.
spawning is common in the highly fecund species, In most shelf waters, the fisheries have influenced
e.g., cod may spawn 10–13 batches of its very small age distributions to the extent that life expectancy is
eggs within a spawning season. significantly reduced.
2440 FISH/Demersal Species of Tropical Climates

0008 Somatic growth patterns also vary widely among most to our understanding of the deep demersal en-
demersal fishes of tropical climates. In fast-growing vironment and the community ecology of fishes there.
species living in strongly seasonal environments, the
overall growth trajectory may show seasonality either Important Species of Demersal Fishes of
because food supply and feeding rates vary, or
Tropical Climates
because the somatic growth is influenced by repro-
ductive activity/gonad growth. Growth rate usually The Clarias
declines significantly when the fish becomes mature,
but never ceases entirely. Attempts have been made to Clarias gariepinus, also known as the African mud 0010

classify species into r- and K-selected types (r and K catfish, exists in the wild but it is also cultivated in
are coefficients expressing rate of reproduction and ponds, cages, and pens and is of great commercial
somatic growth, respectively). The r-selected types importance. This is an omnivorous fish with a prefer-
are those with short life spans but fast growth and ence for a planktonic diet. It also feeds on other types
rather small adult size. They mature at a small size of food items such as insects, insect larvae, pupae,
and young age, and can rapidly take advantage of fish, and fish remains. It also has a propensity for
short and unpredictable favorable environmental being carnivorous. Propagation of these fishes is
conditions by increasing their numbers. The K- widely practiced in the tropics. Peak production of
species, however, tend to invest more energy in som- eggs by the female and hatching of the fertilized eggs
atic growth by growing more slowly and maturing are known to take place during the rainy season. A
later in life, when they are capable of producing many production mean of about 28 000 eggs and a hatching
young. The populations of such species comprise rate of 86.4% have been recorded during the rainy
many age groups, and they are not, like the r-species, season. A low propagation yield at other times has
so vulnerable to repeated recruitment failures due to been attributed to the resting period in the ovarian
low survival of young over a range of years. There are cycle of the animal. Induction of spawning has been
many species that do not readily fit into these classes, carried out with 11-desoxycorticosterone acetate or
and it is unusual for demersal species of tropical carp pituitary suspension. Some of the processed
climates to have the extreme r- or K-strategies seen products made from Clarias include fermented,
among epipelagic fishes and demersal deep-sea fishes, dried, and hot-smoked products. The nutrient con-
respectively. tent of Clarias spp. is given in Table 2.
The Ariomma
Species Distribution Ariomma bondi and A. melanum are the two species. 0011

0009 The vast extent and inaccessibility of the near-bottom The former is found in 200–400 m depth of water; it
oceanic environment have resulted in extremely is silvery and has a big head with small eyes. A.
patchy sampling and arguably therefore a quite re- melanum inhabits 200–600 m of water, is brown in
stricted, and no doubt biased, view of overall demer- color, and has a small head with small eyes. The drift
sal fish distribution and community composition. fish (A. bondi) is a relatively unexploited deep-sea fish
Slope areas in certain regions, especially some parts found in tropical West Africa from Senegal to Gabon
of the North Atlantic, are notable exceptions yet, on and in the western Atlantic from Nova Scotia through
the basis of relative areas, the proportion of the abys- to the Gulf of Mexico. The length range is about
sal floor of the world ocean that has ever been 10–19 cm, with a corresponding average weight of
trawled is so minuscule that any evenly distributed 31.88 g. Size is an important technological property
species represented in collections by more than 20 of a fish, of significance to its fishery and subsequent
specimens in toto is potentially more numerous across utilization. The nutrient content of Ariomma spp. is
the globe than in humanity! The relative uniformity given in Table 2. The length–weight measurement, as
of physical features, such as temperature and salinity well as the weight composition, has been found to
at considerable depth, could allow very widespread
distribution patterns to occur. The detailed know- Table 2 Nutrient content of Ariomma and Clarias spp. tbl0002

ledge of the distribution of most of the demersal Nutrient Composition


species of tropical climates, however, remains
Clarias spp. Ariomma spp.
obscure. Such knowledge is largely derived from
taxonomic and rare zoogeographic studies. Rarely Moisture 70.5–72.0 75.5–77.5
does it come from studies of full assemblages where Protein 18.2–18.6 19.47–20.12
Lipid 6.7–7.1 3.32–3.39
species, ecology, and interrelationships are taken into
Ash 2.6–2.9 2.67–2.71
account. Yet it is these investigations that contribute
FISH/Demersal Species of Tropical Climates 2441

meet the technological requirements of canning. In goals used to be set in terms of maximum sustainable
canned Ariomma, the edible portion increases from yield of a few main species, but a broader and more
51.66 to 59.38% when the whole, headless fish with cautious approach is now being introduced, which
the soft bones and fins are canned, consequently includes consideration of the ecosystem within
increasing the calcium and phosphorus content of which these species are produced and which takes
the product. Canned Ariomma will be cheaper than account of the uncertainty in our assessment of the
canned tuna since it is relatively inexpensive and has a consequences of our activities. The most important
lower processing cost. biological goal is to avoid extinction of species. At a
global level it is evident that economic goals are not
being achieved because the capital and operational
Demersal Fisheries
costs of marine fisheries are about two times higher
0012 Demersal fisheries use a wide variety of fishing than the gross revenue.
methods to catch fish on or close to the seabed. It is There are large numbers of examples of adverse 0017

defined by the gear used, type of fishing activity, and social impacts of changes in fisheries, often caused
varieties of fishes which are caught. A very wide range by the effects of larger, industrial fishing operations
of fishing gear is used in demersal fisheries, the main on the quality of life and standard of living of small-
ones being bottom trawls of different kinds, which scale fishing communities. Biological management of
are dragged along the seabed behind a trawler. Other demersal fisheries has developed mainly from a
methods include seine nets, trammel nets, gill nets, set single-species ‘yield-per-recruit’ model of fish stocks.
nets, baited lines and long lines, temporary or per- The yield is controlled by adjusting the mortality and
manent traps, and barriers. Catches from demersal size of fish that are caught. In many shared inter-
fisheries make up a large proportion of the marine national fisheries, such as those governed by the
harvest used for human consumption and are the European Union, the annual allocation of catch
most valuable component of fisheries on continental quotas is the main instrument of fisheries manage-
shelves throughout the world. ment. This requires costly annual assessment of
0013 Demersal fisheries have been a major source of many fish stocks, which must be added to the operat-
human nutrition and commerce for thousands of ing costs when looking at the economic balance for a
years. Models of papyrus pair trawlers were found fishery.
in Egyptian graves dating back 3000 years. The in- The instruments for limiting the size of fish caught 0018

tensity of fishing activity throughout the world, are mesh sizes, minimum landing sizes, and various
including demersal fisheries, has increased rapidly kinds of escape panels in the fishing gear. These in-
over the past century, with more fishing vessels, struments can be quite effective, particularly where
greater engine power, better fishing gear, and im- catches are dominated by a single species.
proved navigational aids. Many demersal fisheries There are two classes of economic instruments for 0019

are now overexploited and all are in need of careful fisheries management: corrective taxes and subsidies
assessment and management if they are to provide a and property rights. The latter demands less detailed
sustainable harvest. information than the former and may lead to greater
0014 The products of demersal fisheries are mainly used stakeholder participation in the management process,
for human consumption. The species caught tend to because it fosters a sense of ownership.
be relatively large and of high value compared with The management of demersal fisheries will always 0020

typical pelagic species. Demersal fisheries are also be a complex problem, because the marine ecosystem
known as ground-fish fisheries. is complicated and subject to change as the global
0015 The five demersal marine fish with the highest environment changes. Management will always be
average catches over the decade 1990–2000 are based on incomplete information and understanding
Alaska pollock, Atlantic cod, sand eels, blue whiting, and imperfect management tools. A critical step
and Argentine hake (Table 1). All these species spend towards better management would be to monitor
a considerable proportion of their time in midwater. performance in relation to the target objective and
Sand eels spend most of their lives on or in the seabed. provide feedback in order to improve the system.
The vast majority of demersal fisheries take place on One of the changes which have taken place over the 0021

the continental shelves, at depths of less than 200 m. past few years is the adoption of the precautionary
approach. This seeks to evaluate the quality of the
evidence, so that a cautious strategy is adopted when
Management of Demersal Fisheries
the evidence is weak. Whereas in the past such bal-
0016 It is necessary to manage the demersal fisheries for ance of evidence arguments were sometimes applied
biological, social, and economic purposes. Biological in order to avoid taking management action unless
2442 FISH/Pelagic Species of Tropical Climates

the evidence was strong, the presumption now is that groups of typically pelagic fish. Of the total fish catch,
in case of doubt it is the fish stocks rather than the approximately 25 million tonnes are used for reduc-
short-term interests of the fishing industry which tion purposes and it is likely that a very high propor-
should be protected. This is a very significant change tion of this comes from pelagic fish landed in bulk.
in attitude, which gives some grounds for optimism in The production of fish meal from oily fish is around 6
the continuing struggle to achieve sustainable fisher- million tonnes, with an associated 1.4 million tonnes
ies and healthy ecosystems. of oil, implying a conversion ratio of about 3.4:1.
Allowing for approximations, these figures imply
See also: Fish Farming that about 70% of tropical pelagic fish caught are
used for purposes other than direct human consump-
Further Reading tion. In addition to meal and oil, over 1.7 million
tonnes of canned pelagic fish products is produced.
Cochrane KL (2000) Reconciling sustainability, economic The landmass between the tropics of Capricorn 0002
efficiency and equity in fisheries: the one that got away?
and Cancer (32  280 S and N) is considered as the
Fish and Fisheries 1: 3–21.
Cushing DH (1996) Towards a Science of Recruitment in
tropical region. Most of this region, barring some
Fish Populations. Oldendorf/Luhe, Germany: Ecology desert areas in Africa and the Indian subcontinent, is
Institute. considered to be lush with vegetation and animals
Gerking SD (1994) Feeding Ecology of Fish. San Diego: and is characterized by a climate predominantly
Academic Press. determined by annual monsoon rains. From a socio-
Gomes MC (1993) Predictions under Uncertainty. Fish economic point of view, the tropics, in spite of its
Assemblages and Food Webs on the Grand Banks of expected lushness, has some of the poorest and least
Newfoundland. St John’s, Canada: ISER, Memorial developed countries in the world. The great bulk of
University of Newfoundland. mariculture in the tropics takes place on land-based
Gulland JA (1988) Fish Population Dynamics: The Implica- ponds, which draw sea water through natural inlet
tions for Management, 2nd edn. Chichester: John Wiley.
channels, and in shallow bays. Unlike in temperate
Hoines A, Bergstad OA and Albert OT (1995) The food
web of a coastal spawning ground of the herring (Clupea
regions, offshore culture facilities using large floating
harengus L.). In: Skjoldal HR, Hopkins C, Erikstad KE and/or submersible cages are virtually nonexistent. In
and Leinaas HP (eds) Ecology of Fjords and Coastal particular, the almost exponential growth of the
Waters. Amsterdam: Elsevier Science. shrimp culture industry in the last 15 years or so,
Jobling M (1995) Environmental Biology of Fishes. and the adverse environmental influences this devel-
London: Chapman and Hall. opment has had on the coastal environment, and the
Kurlansky M (1997) Cod. A Biography of the Fish that future potential in mariculture development in the
Changed the World. London: Jonathan Cape. tropics as a means of closing the projected gap in
Laevastu T (1996) Exploitable Marine Ecosystem: Their Be- supply and demand for aquatic products, especially
haviour and Management. Oxford: Fishing News Books. for the upper end of the market, by year 2015, have
created a fresh awareness of tropical mariculture.

Pelagic Environment
Pelagic Species of Tropical Microscopic planktonic organisms of the oceans form 0003

Climates the basis of marine food chains and the more complex
food webs. A bucketful of tropical sea water may
G D Khedkar and B V Jadhao, Dr Babasaheb contain several thousand microscopic plants and
Ambedkar Marathwada University, Aurangabad,
animals. Microscopic plant life includes all the
Maharashtra, India
C D Khedkar, College of Dairy Technology, Warud
members of the phytoplankton that provide food for
(Pusad), Maharashtra, India herbivores. The phytoplankton is composed of vari-
N V Chavan, Adarsha College, Hingoli , Maharashtra, ous types of unicellular algae, which range from a
India few microns to several hundred microns in size. The
major groups of algae include the diatoms, which are
Copyright 2003, Elsevier Science Ltd. All Rights Reserved.
the dominant type of phytoplankton. It is essential for
diatoms to remain in the surface waters to carry out
photosynthetic processes, and therefore these algae
Introduction
have developed a variety of adaptations to prevent
0001 Of the total world catch of over 80 million tonnes of sinking. Other common types of phytoplankton in-
fish, roughly 36 million tonnes are accounted for by clude the dinoflagellate algae and coccolithophores.

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