Introduction

One of the most important parameters required for trophic

models of ecosystem is the amount of food ingested (Q) by a
population over a period of time (conventionally a year)
relative to its biomass (B), or Q/B (Polovina 1984;
Christensen and Pauly 1992). This parameter is usually
difficult to obtain and is often replaced by arbitrary guesses.
This paper derives multiple regression models for the
prediction of Q/B from easily obtainable population
parameters. These models also test the hypothesis of no
difference between the Q/B values of freshwater and marine
fishes once shape, size, habitat, temperature and food type
are accounted for.
Materials and methods
Pauly (1986) proposed for the estimation of Q/B the model
Q/B =
tr

tmax
(dW/dt)/K
1(t)
/
tr

tmax
W
t
N
t
dt (1)
where N
t
is the number of fishes of age t, W
t
their mean individual weight,
K
1(t)
(see Eqns 710) their gross food conversion efficiency, t
r
the age at
which fish recruit, and t
max
the maximum age in the population. Eqn (1)
expresses Q relative to the biomass of age-structured populations, but can
be solved for a single representative fish by setting the number of recruits
(N
r
) equal to unity. The key assumptions of Eqn (1) then are (i) that the
population studied is in equilibrium, i.e. that, over all, recruitment
compensates for mortality, as must often be assumed in mass-balance
models (Polovina 1984; Christensen and Pauly 1992); and (ii) that the
fishes in this population grow according to the von Bertalanffy (1934, 1939)
growth formula (VBGF), expressed by the relationship:
W
t
= W

{1 exp [K(t t
0
)]}
b
(2)
where W
t
is the mean predicted weight at age t, W

is the asymptotic
weight, i.e. the mean weight the fish would reach if they were to grow
indefinitely, K the rate (dimension time
1
) at which W

is approached, t
0
the
theoretical age at length zero, and b the exponent of a lengthweight
relationship of the form
W = aL
b
(3)
The parameter b of the lengthweight relationship usually takes values
between 2.5 and 3.5 (Carlander 1950, 1969, 1977). When no b value is
available, growth is assumed to be isometric and b is set at 3, while a is set
equal to the condition factor [c.f.] = W/L
3
(Pauly 1984).
The first derivative of the VBGF expresses the rate of growth, dW/dt
and can be written as
dW/dt = W

3K{1 exp [K(t t

0
)]}
b1
(4)
while the number of individuals in the population from time periods t
1
to t
2
is expressed by
N
2
= N
1
exp [Z(t
2
t
1
)], (5)
whose parameter Z is the coefficient of mortality between t
1
and t
2
. It is
assumed that Eqn (5) reasonably describes the survivorship patterns of
juveniles and young adults, i.e. of those stages contributing most to the
biomass of the population.
The baseline values of Z used here are estimates of natural mortality
(M), obtained from the empirical equation of Pauly (1980), i.e.
logM = 0.0066 0.279 logL

+ 0.65431logK + 04631 logT, (6)

where L

is total length (TL) in cm, K is expressed on an annual basis, T is

temperature in C (see Table 1) and where log, as elsewhere in this
Mar. Freshwater Res., 1998, 49, 44753
10.1071/MF98015 1323-1650/98/050447
Predicting food consumption of fish populations as functions of mortality,
food type, morphometrics, temperature and salinity
Maria Lourdes D. Palomares
A
and Daniel Pauly
B
A
International Center for Living Aquatic Resources Management, MC PO Box 2631,
Makati City, 0718 Metro Manila, Philippines.
B
Fisheries Centre, 2204 Main Mall, University of British Columbia, Vancouver, BC Canada, V6T 1Z4.
Abstract. A large data set of relative food-consumption estimates (Q/B) of marine and freshwater fish
populations (n = 108 populations, 38 species) is documented and used to derive a predictive model for Q/B,
using asymptotic weight, habitat temperature, a morphological variable and food type as independent
variables. Salinity is shown to have no effect on Q/B in fish well adapted to fresh or salt water (other things
being equal), while mortality (Z), has a strong, positive effect on Q/B and on gross food-conversion
efficiency (defined by GE = Z/(Q/B)), by affecting the ratio of small:large fish. The empirical models thus
derived should be useful for parameterization of trophic models of ecosystems and similar applications.
CSIRO 1998
Fig. 1. Aspect ratio of the caudal fin for two generic fish, a very active
tuna (A = 6.7) and a less active grouper (A = 0.7).
Palomares, M.L.D. and D. Pauly. 1998. Predicting food consumption of fish populations as functions of mortality, food type,
morphometrics, temperature and salinity. Marine and Freshwater Research 49: 447-453.
Maria Lourdes D. Palomares and Daniel Pauly 448
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5
1
Maria Lourdes D. Palomares and Daniel Pauly 450
contribution, refers to log
10
. To investigate the effect of mortality on Q/B,
and to derive predictive models of Q/B taking explicit account of different
mortalities, values of Q/B were calculated, via Eqn (1), for mortalities equal
to 0.5M, M, 2M and 4M.
The food conversion efficiency (K
1
) used in Eqn (1) is defined by
K
1
= growth increment/food ingested (7)
for any time period (Ivlev 1945). Pauly (1986) showed that K
1
depends,
among other things, on the individual weight of fish (W) in a manner that
can be summarized by
K
1
= 1 (W/W

(8)
where is an exponent estimated as the slope of a linear regression of log
1K
1
v. logW. (Pauly 1986). Note that Eqn (8) predicts a K
1
value of zero
at W as should be the case. On the other hand, a value of K
1
= 0 is
predicted for W = 0, which is not realistic. However, this applies to
weights that do not contribute more than a minuscule fraction of a
populations biomass.
Combining Eqn (7) with the VBGF and simplifying allows expression
of K
1
and dW/dt (see Eqns (4) and (7)) as a function of age t through
K
1(t)
= 1 {1 exp [K(t t
0
)]}
b
(9)
The application of Eqn (1) is not limited to estimates of K
1
obtained
from feeding experiments with captive fish. Daily ration (R
d
) estimates can
also be used by first converting them to K
1
through
K
1
= (dW/dt) / R
d
(10)
as is implied from the very definition of K
1
(see Eqn 7).
Daily ration estimates may be obtained from: analysis of 24 h cycles of
stomach contents data obtained from field studies (Jarre et al. 1991), from
metabolic studies (Winberg 1956; Mann 1978), and from nitrogen and/or
energy budget studies (Ivlev 1945; Gerking 1962, 1978).
The methods and equations presented above were applied to estimates of
K
1
and R
d
in 108 freshwater and marine populations of fishes and were used
to estimate Q/B values. Then, multiple regression models were constructed
which included various parameters earlier hypothesized to affect Q/B, i.e.
the VBGF parameter W

(in g), with a slope expected to be negative

(Palomares and Pauly 1989; Pauly 1989); the mean annual temperature of
the water body, expressed as T = 1000/Kelvin (Kelvin = C + 273.15). A
positive relationship between temperature and Q/B is expected. However,
given the transformation used here, a negative slope should be estimated
(Regier et al. 1990);
The aspect ratio of the caudal fin which is a measure A of the swimming
and metabolic activity of the fish expressed as
A = h
2
/s (11)
where A is the aspect ratio, h the height of the caudal fin and s the surface area
of the caudal fin, extending to the narrowest part of the caudal peduncle
(Fig. 1); a positive slope is expected (Pauly 1989); two morphometric ratios
as indices of the body form of a fish, i.e. D = standard length / body depth,
and P = caudal peduncle height / body depth. High Q/B values were
previously hypothesized to be associated with intermediate values of D, while
a negative slope was hypothesized for P (Pauly 1989); the types of food
consumed, i.e. h for herbivores (h = 1, d = 0 and p = 0; positive slope); d for
detritivores (d = 1, h = 0 and p = 0; positive slope); and p for cultured fish fed
with commercial pellets (p = 1, h = 0 and d = 0; no prior hypothesis
concerning slope); carnivores are identified by default (h = 0, d = 0 and p = 0)
(Palomares 1991); and the salinity of the water body, i.e. S = 1 for marine or
brackishwater and S = 0 for freshwater (no hypothesis concerning slope).
Estimates of Q/B obtained from the data in Table 1 were recalculated for
values of Z derived by multiplying M by 0.5, 2 or 4. These were added,
along with the other (unchanged) predictor variables, to the file containing
the original Q/B values and related predictor variables, and in which Z = M
pertained. Then, multiple regression models were calculated from this
artificially expanded data set by using either Z or the M-multiplication
factor (f) as an additional predictor variable. The regression statistics in
such cases are meaningless, because Q/B is statistically related to Z (and f)
through the incorporation of Eqn (5) into (1), and because the expanded
data set consists essentially of replicates of the data set in Table 1.
Results and Discussion
Table 1 presents the Q/B and related statistics assembled
here for 108 populations of fish, distributed over 65 species
in 25 families of teleosts, of which 40 occur in marine and
68 in freshwater. The exploratory multiple regression model,
including all hypothesized variables, is documented in Table
2; the slopes associated with depth ratio, peduncle depth,
pellets and salinity have extremely large standard errors,
indicating that they do not have significant relationships
with Q/B. The slope associated with aspect ratio, on the
other hand, is a borderline case, notwithstanding a formal
test. Given this, and the fact that it is the morphometric
variable with the lowest standard error, it was kept in Eqn
(12), the second model estimated
logQ/B = 7.964 0.204logW

1.965T+ 0.083A (12)

+ 0.532h + 0.398d
whichexcept for Aincludes only terms significant at
P<0.001 (Table 2). All the signs in Eqn (12) are as
previously hypothesized. Fig. 2a shows how the predicted
values of Q/B correlate with the observed values, and Figs
2b2d show that the residuals do not form patterns with
regard to any of the re-expressed continuous variables. The
residuals are normally distributed (Fig. 2e), thus suggesting
that the key assumptions of linear regression were met.
Table 2. Statistics of exploratory multiple linear regression, and of Eqns
(1214), linking Q/B and predictor variables (n = 108)
(Intercepts in parenthesis)
Exploratory model Eqn (12) Eqn (13) Eqn (14)
estimate s.e. estimate s.e. estimate estimate
Intercept (6.972) (0.340) (7.964) (0.339) (8.056) (5.847)
Slope
Factor (log) 0.300
Z (log; year
1
) 0.280
W

(log; g) 0.209 0.036 0.204 0.033 0.201 0.152

T(1000/K) 1.729 0.476 1.965 0.406 1. 989 1.360
Aspect ratio 0.071 0.046 0.083 0.044 0.081 0.062
Depth ratio 0.291 0.417
Peduncle depth 0.372 0.330
Herbivore
A
0.517 0.112 0.532 0.104 0.522 0.510
Pellets
A,B
0.060 0.095
Detritivore
A
0.352 0.153 0.398 0.144 0.393 0.390
Salinity 0.058 0.083
R
2
0.530 0.516
df 98 102
A
Dummy variable.
B
Variable that identifies (aquaculture) populations for which only the amount
of food given is known and not the amount actually consumed by the fish.
451 Predicting food consumption of fish populations
Mitchell (1997) argued that the approach documented in
Fig. 2 to validate a predictive model is inappropriate and
suggested the use of the approach in Fig. 3 instead. This also
suggests that Eqn (12) meets the expectations for a model
such as sought here.
[Note also that the circularity affecting Z (and f; see above)
is avoided in the case of W

, although this parameter enters

Eqn (12) both as independent variable and as factor in Eqns
(2) and (4), used to estimate Q/B, the dependent variable.
This is so because the values of W

cancel out when (2)

Fig. 2. Key features of the predictive model in Eqn (12). (a) Plot of log predicted v. log observed values; note relatively tight fit
(r = 0.718, df 106). (b) Residuals (log predicted log observed values) v. logW

; note absence of pattern, though W

values between
100 and 10 000 g do predominate. (c) Residuals v. transformed temperature (1000/Kelvin); note faint vertical patterns due to preferences
for round values (in C). (d) Residuals v. caudal-fin-aspect ratio; scarcity of high values (for tunas) explains the non-significant effect
of this biologically important variable. (e) Frequency distribution of residual Q/B values; note normality of distribution.
P
r
e
d
i
c
t
e
d

Q
/
B
(
l
o
g
;

y
e
a
r

1
)
Observed Q/B (log; year
1
)
R
e
s
i
d
u
a
l

Q
/
B
(
l
o
g
;

y
e
a
r

1
)
F
r
e
q
u
e
n
c
y
(c) (b)
(a)
Temperature (T; 1000/K)
Caudal fin aspect ratio (A)
Asymptotic weight (log; g)
Residual Q/B (log; year
1
)
(d) (e)
R
e
s
i
d
u
a
l

Q
/
B
(
l
o
g
;

y
e
a
r

1
)
R
e
s
i
d
u
a
l

Q
/
B
(
l
o
g
;

y
e
a
r

1
)
Maria Lourdes D. Palomares and Daniel Pauly 452
and (4) are inserted into (1). The other independent variables
of Eqns (12)(16) are completely unrelated to Q/B.]
Thus validated, the statistics in Table 2 and Figs 2 and 3
lead to three major conclusions:
(1) Q/B can be predicted from population parameters that are
relatively easy to estimate;
(2) morphometric variables for body depth and for caudal
peduncle height, found by Pauly (1989) to have a
significant effect on Q/B, have no relationship with Q/B
once food type and temperature are properly accounted
for; and
(3) for fish well adapted to fresh or salt water, salinity, other
things being equal, does not have any significant effect on
Q/B, contrary to a widespread, if often unstated, belief.
Re-expressing Eqn (12) in terms of path coefficients
(Blalock 1972), i.e. of partial slopes standardized by the
standard deviations of the variates, leads to the following
values: logW

= 0.455; h = 0.354; T = 0.340; d = 0.193

and A = 0.141. Thus, the effect of asymptotic weight on Q/B
is about three times as strong as the effect of aspect ratio,
with the other variables taking intermediate ranks.
The addition to our original data set (n = 108) to cover
Q/B values recomputed for multiplication factors (f) of M
equal to 0.5, 2 and 4 led to Eqn (13):
log Q/B = 8.056 + 0.300logf 0.201logW

+ 1.989T (13)
+ 0.081A + 0.522h + 0.393d
where all variables are defined as in Eqn (12), except for f,
which has a positive slope, as expected. This model can be
used to predict Q/B of exploited stock whose exact Z value is
unknown, but for which total mortality can be approximated,
based on natural mortality times a factor. Eqn (14) is supplied
for cases where the exact value of Z is known:
log Q/B = 5.847 + 0.280 logZ 0.152 logW

1.360T (14)
+ 0.062A + 0.510h + 0.390d
where Z is equal to M + F, the latter term representing
fishing mortality.
Allen (1971) has shown that for a variety of expressions
for growth and mortality (including Eqns 2 and 5) the
instantaneous rate of total mortality Z is equal to production/
biomass ratio (P/B), a key input in trophic models (Christensen
and Pauly 1992).
In analogy to K
1
, which expresses the food conversion
efficiency of (a group of) animals of a given size, a gross
conversion efficiency (GE) can be defined, for an age-
structured population as
[GE] = (P/B) / (Q/B) (15)
Thus, given the identity between Z and P/B, we can derive
from Eqn (14) the model
log[GE] = 5.847 + 0.720logZ + 0.152logW

+ 1.360T (16)
0.062A 0.510h 0.390d
which predicts GE as a function of Z, asymptotic weight,
temperature and food-related parameters.
Here, as for Eqns (12)(14), values of A = 1 may be used
for fish which do not use their caudal fin as main organ of
propulsion, and which will therefore tend to have low meta-
bolic rates (Pauly 1989; see also Fig. 1 and Fig. 2d).
These equations will be useful for parameterization of
trophic models. A previous model (Palomares and Pauly
1989) was widely used for such purposes (see, e.g., contrib-
utions in Christensen and Pauly 1993), although it did not
include as many cases and as many variables as the models
presented here.
Acknowledgments
We acknowledge Mr Francisco B. Torres Jr for his help with
spreadsheet computations and Dr Villy Christensen for his
useful comments on the manuscript. M.L.D.P also thanks
Professor Jacques Moreau for his supervision of the work done
at the Ecole Nationale Suprieure Agronomique de Toulouse,
France. This paper is ICLARM Contribution No. 1414.
References
Allen, K. R. (1971). Relation between production and biomass. Journal of
the Fisheries Research Board of Canada 28, 157381.
Bertalanffy, L. von. (1934). Untersuchungen ber die Gesetzlichkeiten des
Wachstums I. Roux Archiv fr Entwicklungsmechanik 131, 61352.
Bertalanffy, L. von. (1939). A quantitative theory of organic growth
(inquiries on growth laws II). Human Biology 10, 181213.
Blalock, H. M., Jr (1972). Social Statistics. (McGraw-Hill: New York.)
583 pp.
Carlander, K. D. (1950). Handbook of Freshwater Fishery Biology.
(Dubuque: Iowa; Wm. C. Brown: Ames.)
Carlander, K. D. (1969). Handbook of Freshwater Fishery Biology. Vol.
I. (Iowa State University Press: Ames.)
Carlander, K. D. (1977). Handbook of Freshwater Fishery Biology. Vol.
II. (Iowa State University Press: Ames.)
Christensen, V., and Pauly, D. (1992). The ECOPATH IIa software for
balancing steady-state ecosystem models and calculating network
characteristics. Ecological Modelling 61, 16985.
Fig. 3. Plots of residuals (log predicted log observed values) v.
predicted values. Note absence of pattern, which validates Eqn (12) by the
criterion of Mitchell (1997).
Predicted Q/B (log; year
1
)
R
e
s
i
d
u
a
l

Q
/
B
(
l
o
g
;

y
e
a
r

1
)
453 Predicting food consumption of fish populations
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Manuscript received 3 February 1998; revised and accepted 28 June 1998