Fish swim in a surprising number of ways |
Immersion in water is the closest approximation to
weightlessness. Moreover, teleosts, through their invention of the gas bladder,
have learned to escape the drag of gravity with the least energy expenditure.
The downside of being under water is the fact that any movement has to push
aside water, a particularly dense and heavy medium. The physics and physiology
of fish swimming are summarized in Blake (1983) and Webb (1984) and the most
recent comprehensive review in this field is that of Videler (1993).
This section describes the manner in which fish swim, something
they do in a surprising number of ways; also, available records on the swimming
speed of fish are presented.
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We follow Lindsey's classification |
The classification of the swimming modes of fish adopted here
stems from Lindsey (1978), who reviewed much of the earlier literature on this
topic. This classification consists of two levels, the first (Roman numerals)
describing what may be called the swimming ‘type’, the second (bullets)
describing the swimming ‘mode’ proper,
viz.
Movements of body and/or caudal fin:
Anguilliform;
Subcarangiform;
Carangiform;
Thunniform; and
Ostraciiform.
Undulation of median or pectoral fins:
Amiiform;
Gymnotiform;
Balistiform;
Rajiform; and
Diodontiform.
Oscillations of median or pectoral fins:
Tetraodontiform;
Labriform.
The modes in (I) imply a gradual transition from undulation of
the entire body (including the trunk) being used for propulsion (anguilliform
mode) to propulsive forces being generated only by the oscillating caudal fin
(thunniform and ostraciiform modes).
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A gradual transition from undulation to oscillation |
The ranking of the modes in I and II to III further implies a
gradual transition from undulations to oscillations as the movements generating
the major propulsive force, and thence the above scheme can also be represented
as a graph, with a trunk-to-caudal-fin axis as ordinate and an
undulation-to-oscillation axis as abscissa (see Fig. 1 in Lindsey 1978).
This implies that the assignment of a swimming mode to a given
fish species will always contain a subjective element, even if we ignore the
fact that fish may have two swimming modes (we consider here only the dominant
mode, e.g., labriform in parrotfish, Family Scaridae, even if parrotfish revert
to the subcarangiform mode when escaping from presumed danger (authors’ pers.
field obs.)).
Thus, the entries of this table (i.e., the choices of the types
and modes listed above) may be revised from time to time as a result of our
familiarization with this topic and its literature. This will not, however,
affect the first set of entries, based on Lindsey’s examples [Species explicitly
assigned to a certain swimming mode by Lindsey (1978) in both the
MainRef. and Ref. fields] and their obvious extensions (e.g., from
Anguilla anguilla to all Anguillidae, and thence to all Anguilliformes).
Note also that this table presently pertains only to juvenile
and adult fishes. Fish larvae¾ for obvious reasons¾ have a limited repertoire of swimming types and modes.
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The caudal fin as life-style indicator |
The aspect ratio of the caudal fin of a species closely
correlates with its average level of activity (Pauly 1989). The Aspect ratio
(A) of the caudal fin is calculated from:
A = h2 / s
where h is the height of the caudal fin and s its
surface area (Fig. 52). The proximal border of the caudal fin surface is defined
as a straight vertical line through the narrowest portion of the caudal
peduncle, i.e., the portion of the peduncle surrounded by the fin is considered
part of the caudal fin area (see Fig. 52).
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Fig. 52. Aspect ratio (A = h2/s, h = height of the
caudal fin; s = surface area of fin) of a pelagic fish (A = 7.5) and a bottom
dweller (B = 0.6). Note the correspondence between aspect ratios and modes of
life.
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Note that this definition of A differs slightly from
that presented in Pauly (1989), where only the caudal fin proper was used for
calculating s.
A reference may be given to document an aspect ratio; when none
is given, the aspect ratio was calculated, by planimetry, from a fish picture in
FishBase or another readily available source.
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Status |
The SWIMMING table contains swimming type and mode for over
2,700 species.
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Sustained speed and burst speed |
The SPEED table contains 255 records with maximum swimming
speeds for 80 species. The information was extracted from over 50 references
such as Bainbridge (1958, 1960), and Webb (1971) and compilations such as
Sambilay (1990). An effort was made to distinguish between ‘sustained’ (i.e.,
maintained for more than 3 minutes), and ‘burst’ swimming (maintained for a few
seconds only) (see Fig. 53), as well as other swimming modes (Hammer 1995).
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Fields |
The SPEED table consists of the following fields:
Length: This field pertains to the length of fish in cm as
stated in the publication. The type of length measurement used consists of the
following choices: SL (Standard Length); FL (Fork Length); TL (Total Length); BL
(for the term ‘body length’, stated in the publication but without the type of
length measurement being indicated).
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Fig. 53. Relationship between swimming speed and body length of
fishes. Note that burst speed is about 10 times higher than sustained speed.
Note also that no fish appears to swim at speeds below Weihs’ (1973) line of
minimum energy expenditure per distance covered, whose slope (1 : 1, implying
strict proportionality of speed and length), however, is the same as that of the
observations in this figure.
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The Swimming speed gives the speed of the fish per
second as reported in the source. A choice field is provided for the type of
length measurement used. The choices are the same as above (SL, FL, TL, BL) and
m/s (for meter per second). A calculated Speed field in meter per second
is also provided (see Derived values).
Mode: This field describes the mode of swimming as:
sustained (swimming at this speed for a prolonged time); burst (a maximum
swimming speed which can be maintained for less than a minute only);
other.
Comment: States the length or weight measurement, or mode
of swimming if not in the choice list. Any transformation of length type should
also be stated here.
Derived values: Gives the transformation values from
different units of speed to standard length per second (SL/s) and to meter per
second (m/s). This makes it possible to compare the swimming performance of
fishes with forked, rounded or other shapes of caudal fin. Where available, the
a and b values used to transform fork length or total length to
standard length are given as used in:
SL = a + b × L
where L is the reported fork or total length. If the
transformation is based on one measurement only, hopefully a typical adult, the
intercept a is set to 0 (see also the ‘LENGTH-LENGTH table’, this
vol.).
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How to get there |
You get to the SWIMMING and SPEED tables by clicking on the
Biology button in the SPECIES window, the Morphology and
physiology button in the BIOLOGY window, and either the Swim. type or
Swim. speed buttons in the next window.
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Internet |
On the Internet, you get to the SPEED table by clicking on the
Speed link in the ‘More information’ section of the ‘Species Summary’
page. You can create a list of all species with available data by selecting the
Swim. Speed radio button in the ‘Information by Topic’ section of the
‘Search FishBase’ page. As of December 2000, information on swimming type was
not yet available on the Internet.
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References |
Bainbridge, R. 1958. The speed of swimming of fish as related
to the size and to the frequency and amplitude of the tail beat. J. Exp. Biol.
35(1):109-133.
Bainbridge, R. 1960. Speed and stamina in three fish. J. Exp.
Biol. 37(1):129-153.
Blake, R.W. 1983. Functional design and burst-and-coast
swimming in fishes. Can. J. Zool. 61:2491-2494.
Hammer, C. 1995. Fatigue and exercise tests with fish. Comp.
Biochem. Phsyiol. 112A(1):1-20.
Lindsey, C.C. 1978. Form, function, and locomotory habits in
fish, p. 1-100. In W.S. Hoar and D.J. Randall (eds.) Fish physiology.
Vol. II. Academic Press, New York.
Pauly, D. 1989. A simple index of metabolic level in fishes.
Fishbyte 7(1):22.
Sambilay, V.C., Jr. 1990. Interrelationships between swimming
speed, caudal fin aspect ratio and body length of fishes. Fishbyte
8(3):16-20.
Videler, J.J. 1993. Fish swimming. Chapman and Hall, London.
260 p.
Webb, P.W. 1971. The swimming energetics of trout. Thrust and
power output at cruising speed. J. Exp. Biol. 55:489-520.
Webb, P.W. 1984. Form and function in fish swimming. Sci. Am.
251:58-68.
Weihs, D. 1973. Optimal fish cruising speed. Nature 245 (Sept.
7):48-50.
Rainer Froese, Armi Torres, Crispina Binohlan and Daniel
Pauly
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