Hydrobiologia 237 : 61-66, 1992 .

© 1992 Kluwer Academic Publishers . Printed in Belgium . 61

Effects of angling on population structure of brown trout, Salmo

trutta L ., in mountain streams of Northern Spain

F . Brava, A. G . Nicieza & M . M . Toledo

Departamento de Biologia de Organismos y Sistemas (Zoologia), Universidad de Oviedo, 33005 Oviedo,
Spain

Received 5 December 1990 ; in revised form 16 July 1991 ; accepted 22 August 1991

Key words : angling exploitation, trout fishery, age distribution, size distribution, density

Abstract

Effects of angling exploitation on brown trout populations were assessed by comparing fished sections
with close ones unfished for at least 20 years, in mountain streams of Asturias (Northern Spain) . Both
the fish size and age structure significantly differed among sections in the expected direction according
to their exploitation status . The main effects were a significant decrease in age structure complexity
(diversity), life span, and percent individuals above the legal limit size in the exploited stocks versus the
unexploited ones . Trout above the minimum length limit for fishing (18 cm) averaged 19 .47% of the fish
caught in the unfished sections (sd = 4.01 ; n = 5), and 4 .72% (sd = 3 .46 ; n = 4) in those subjected to
angling . Furthermore, fish older than 4 years represented 39.84% (sd = 8 .53) and 1 .19% (sd = 1 .60) of
the catch, respectively . Effects on recruitment (density of young fishes) and growth rates (length at age
1 + to 3 + ) were not absolutely consistent, though maximum values were associated with fished sections .

Introduction and fecundity (Healey, 1978, 1980 ; Fraser et al.,

Most trout populations in lakes and streams are
subject to some angling exploitation, which ac-
counts for a significant mortality of the oldest
age groups (Shetter, 1969 ; Avery & Hunt, 1981 ;
Swales & Fish, 1986) . In extreme cases, angling
mortality can so reduce the spawning stock that
recruitment becomes insufficient to sustain the
fishery without stocking (Skurdal et al., 1989).
The loss of reproductive potential may be espe-
cially harmful in slow growing, late maturing
populations of brown trout, such as those found
at high altitudes in the Cantabrian Mountains
(Garcia & Braila, 1988) . Moreover, alteration in
population structure caused by fishing can also
affect patterns of habitat use, activity, growth rates
1987 ; Donald & Alger, 1989) . Despite the obvi-
ous need to understand the responses of salmo-
nid populations to exploitation, such studies are
relatively scarce (Donald & Alger, 1989) .
Our objective was to assess the impact of an-
gling on brown trout populations in mountain
streams of Asturias (Northern Spain), by exam-
ining differences in age structure, size structure,
and density between closely related stocks, some
of them unexploited for almost 20 years and
others subject to sport fishing .
Material and methods
Seven sections of three mountain streams (eleva-
tion from 500 to 890 m) in the high basin of the
62
43°
10'
43 0
00'
10 Km
6 ° 45'
Fig. 1 . Study area showing the sampled sections and location of the Muniellos Biological Reservation .
river Narcea (Fig. 1) were electrofished for brown
trout (Salmo trutta L .) . Three of these sections are
within the Biological Reservation of Muniellos
and have been closed to fishing since 1972 . The
remaining four sections, including one on the river
Muniellos downstream of the Biological Reser-
vation, are fished by recreational anglers from
early May to the second half of August . There is
a bag limit of 16 trout per day and a minimum
length limit of 18 cm . Resident brown trout was
the only fish species present .
All the sampled sections (first or second order
streams) are predominantly riffle zones (mean
depth for sections varied from 16 .2 to 31 .8 cm ;
overall mean ± 1 sd = 25 .4 ± 5.8 cm), and are quite
homogeneous with respect to habitat structure .
The two upper sections on the river Muniellos
(Zreizal & Banzao) are located above an unpass-
able dam, so immigration from downstream do
not exists at these points . Area of the sampled
sections averaged 271 m2 (sd = 97 ; range : 172-
448) . Population sizes were estimated using
Zippin's (1956) removal method . The estimates
F BIOLOGICAL RESERVATION
1 - ZREIZAL
2 - BANZAO
3 - TABLIZAS
4 - MOAL
5 - GEDREZ
6 - COTO
7 - NAVIEGO
.-\
6° 00'
BISCAY BAY
6 ° 30' 6-15'
were made 1 to 3 times in each section from
November 1988 to August 1989 (Table 2) . The
fork length of all fish caught was measured to the
nearest 1 mm. Age was determined from scale
samples taken from almost all the specimens
larger than 11 cm and from some below this size .
Age of the remaining fish was assigned by exam-
ination of length frequency distributions (Fig. 2) .
Age structure complexity was assessed for each
population by means of Pielou's (1969) diversity
index : D = 1 - E p?, where p, is the proportion
of fish belonging to the ith age class .
Growth was determined by using age specific
length distributions for the different sections . The
data were analysed by applying ANOVA and
multiple comparison tests (Scheffe procedure) .
Logarithmic transformations were used to nor-
malize length data . The Mann-Whitney U test
was used to compare proportions above selected
length and age limits between exploited and un-
exploited sections . The rejection level for all tests
was set at p = 0.05 .


63

2 + -- --•
3 +
1

3 +

5 _- 4 +

0 -

0 5 10 15 20 25 30

FORK LENGTH (cm)

.8+
7 + 1 - ZREIZAL-AUGUST 4 - MOAL-AUGUST

3C 2 - BANZAO-MARCH 5 - GEDREZ-AUOUST
3A - TABLIZAS-NOVEMBER 6 - COTO-AUGUST

0 3B - TABLIZAS-MARCH 7 - NAVIEOO-AUGUST
3C - TABLIZAS-AUGUST
FORK LENGTH (cm)
Fig . 2 . Length frequency distribution for brown trout caught from November 1988 to August 1989 at different sections in the high
basin of the Narcea river . Sections (or dates within section) labelled 1, 2 and 3 (A, B, C) were unexploited for nearly 20 years,
and those labelled 4, 5, 6 and 7 were subjected to angling exploitation . Ages determined by scale reading were shown over each
histogram, where black boxes indicate trout above the size limit for fishing .
64

Results index, showed significantly higher values in un-

By August, mean fork length of brown trout
caught in different river sections ranged from 8 .2
to 10 .7 cm for age group 1 +, 11 .6 to 17 .5 for age
group 2 + and 14 .5 to 19 .7 for age 3 + . By this
time, the size distributions differed among sec-
tions at each age from 1 + to 3 + (ANOVA,
p<0 .001 in all cases) . Several significant differ-
ences in fork length were evident between sec-
tions, especially for Coto and Naviego (two fished
sections), which had more rapid growth than the
remaining sections (Table 1) . The sequence of
localities ranked in order of increasing fork length
was the same for ages 1 +, and 2 +, and almost
the same for 3 + . Trout in age-group 0 + were
omitted for this analysis because of their small
number in some sections and the possible size
selection by electrofishing at this age, and those
older than 4 years because they were not present
in all sections and were likely to be size-biased by
angling harvest .
Populations from unexploited sections showed
a more complex age structure and longer life span
than those that were fished (Table 2 ; Fig. 2) . Age
structure diversity, as measured by Pielou's (1969)
exploited sections (Table 2 ; Man-Whitney U test,
p<0 .01) . The minimum size limit (18 cm) was
reached at age 3 + at Coto and Naviego, but not
before age 4 + or 5 + in the remaining sections .
In the unfished sections brown trout above 18 cm
comprised from 15 .3% to 23 .5% of the total
number of fish caught (Table 1 ; mean = 19.5% ;
sd = 4 .0 ; n=5), but constituted only from 0 to
8.3% in the exploited sections (mean = 4 .7 % ;
sd = 3 .5 ; n = 4) . Similarly, the percentage of indi-
viduals of age equal or older than 4 years aver-
aged 39 .8 % (sd = 8.5 ; n = 5) in the unfished sec-
tions and 1 .2 % (sd = 1 .6 ; n = 4) in those subjected
to angling pressure . The differences in age and
size of fish from exploited and unexploited sec-
tions were both significant (Mann-Whitney U
test ; p<0 .01) .
Trout density ranged from 16 .1 to 40 .0
fish 100 m -2 (Table 1), and there were no con-
sistent differences between sections. However, the
values from unexploited sections varied less than
those from sections subjected to fishing . When
only the density of trout below the fishing size was
considered, two exploited stocks (Naviego and
Moal) showed maximum values, although these

Table 1 . Mean fork length (cm) for age groups 1 + to 3 + in sections sampled at August . F values corresponding to analysis of
variance for sections within age groups, and results of Scheffe multiple comparison test are also shown . For age group 3 +, Coto
and Naviego sections were pooled because of the small sample size .

Sections Age classes

1+ 2+ 3+

x sd n z sd n x sd n

Zreizal (Z) 8 .23 0 .51 7 11 .60 0 .58 8 14.47 0 .93 10

Tablizas (T) 9 .49 0 .99 20 13 .67 0.39 7 15 .77 1 .18 8
Moal (M) 9 .05 0 .73 20 12 .75 0 .88 13 15 .56 1 .44 21
Gedrez (G) 9 .63 1 .25 14 14 .78 0.83 6 15 .71 0 .94 6
Coto (C) 10 .52 1 .07 11 16 .10 1 .16 11 19 .73 0 .66 3
Naviego (N) 10 .75 1 .45 30 17 .50 1 .88 4 19 .25 - 2

F (a) 9 .77*** 38 .44*** 12 .85***

Scheffe (b) (C, N) (Z, M, T, G) (Z, M) (M, T) (T, G) (Z, M, T, G)
(T, G, C) (G, C) (C, N)

(a) ***p<0 .001 .

(b) Sections without significant differences among them (p>0 .05) are included in the same parentheses .
 65

Table 2 . Main structural characteristics for brown trout stocks sampled at different sections in the upper reaches of the Narcea
basin .

Section Month No Percent fish Percent fish Age structure Density (fish/ 100 m2 ± 95 % c .l .)
>18cm 4 years diversity
Whole population Fish below 18 cm

Zreizal August 43 23 .3 39 .5 0 .813 36.2±24 .2 29 .2±25 .2

Banzao March 85 15 .3 43 .5 0 .817 30 .4+ 10 .9 24 .1+7 .4
° Tablizas November 96 19 .8 43 .8 0 .776 28 .9+2 .0 23 .0+ 1 .5
a
Tablizas March 68 23 .5 47 .1 0 .826 17 .6+3 .4 16 .1+7 .7
Tablizas August 71 15 .5 25 .4 0 .810 36.3+8 .6 32 .8+12 .6

ti Moal August 59 5 .1 3 .4 0 .707 33 .5+8 .2 32 .9+9 .8

.o Gedrez August 31 0 .0 0 .0 0 .695 19 .5+3 .1 19 .5+3 .1
Coto August 36 8 .3 0 .0 0 .712 16.1+1 .6 14 .6+ 1 .3
w
Naviego August 73 5 .5 1 .4 0 .584 40 .0+7 .5 37 .8+7 .3

values were close to those recorded for sections
inside the Reservation in August, and another
two fished sections (Gedrez and Coto) showed
the minimum values in the same period (Table 2) .
Discussion
Age and length-frequency distributions of fish
caught in all river sections indicates slow-growing
populations (Fig . 2 ; Table 1), similar to those re-
ported for streams in northern Europe (Jonsson,
1977 ; Jonsson & Sandlund, 1979; Mortensen,
1977) . Samples from Tablizas show that growth
occurs mainly from March to August, and this
agrees with findings in some other brown trout
populations (Jonsson, 1977 ; Scarnecchia, 1983) .
Brown trout populations from unfished and fished
sections in the high basin of the Narcea river
showed significant differences in age and size
structure which were consistent with their exploi-
tation status . Although slow growing populations
may exhibit delayed maturity and longer life span
than faster growing populations (Jonsson, 1977 ;
Donald & Alger, 1986), differences in growth in-
tensity at the Narcea river basin did not explain
all of the differences in population structure. In
fact, brown trout from two neighbouring sections
of the river Muniellos, Tablizas and Moal, the
first closed to fishing and the second, located
some 400 m downstream, open to fishing, showed
differences in size/age structure, but not in
growth patterns as revealed by length at age for
groups 1 + to 3 + . However, in general exploited
sections (Naviego, Coto, Gedrez) exhibited faster
growth than the unexploited ones, in agreement
with the findings of several authors (e .g ., Healey,
1980 ; Donald & Alger, 1989) .
Our results did not show a distinct reduction in
fish density in the exploited sections, especially
when considering fish below 18 cm . This implies
that fishing pressure and angling regulations (fish-
ing period, size and bag limitation) allow repro-
duction of sufficient individuals to maintain the
population in these sections . Furthermore, this is
a common outcome of exploitation (Healey, 1980 ;
Donald & Alger, 1989), because removal of old
fish can increase juvenile survivorship . Old fish
might relegate the young ones to marginal habi-
tats or restrict their activity times by interference
or by predatory dissuasion, with subsequent in-
crease in mortality (Healey, 1980 ; Werner et al.,
1983 ; Fraser et al., 1987). Large, old fish fre-
quently prey on the young ones (Lopez Alvarez,
1984 ; see also Suarez et al., 1988, for brown trout
in the Narcea river basin), so that cannibalism
may be an important direct mechanism in regu-
lating the abundance of young fish, as reported
for other salmonid populations (Ricker, 1950 ;
Johnson, 1976) .
A common feature of trout populations sub-
jected to sport fishery, even under moderate
66
angling pressure, is the absence of old fish be-
cause cropping through successive fishing periods
prevents them from attaining their potential lon-
gevity (Avery & Hunt, 1981 ; Swales & Fish, 1986 ;
McDonald & Hershey, 1989) . As an example, the
largest fish caught in the unfished sections of the
Muniellos river was 11 years old, and this fish
attained the legal size limit after its third growth
season (back-calculated length at this age was
18 .9 cm) . Trouts living in fished sections hardly
attained this age because it would survive 8 fish-
ing seasons .
The absence of large individuals may have
different effects on a fish population, some of
them advantageous for fishing exploitation, as in-
creased growth rates or juvenile density (Healey,
1980 ; Donald & Alger, 1989) . But fish predation
also acts in multiple, sometimes subtle ways on
habitat selection, size structure, feeding intensity,
growth or age at maturity (Werner et al., 1983 ;
Fraser et al., 1987 ; Huntingford et al ., 1988), so
that survival of some old individuals would be
necessary to allow the development of population
interactions and to maintain a quality fishery .
Acknowledgements
Funding for this study was provided by the En-
vironmental Agency (Principado de Asturias) and
FICYT .
References
Avery, E . L . & R. L . Hunt, 1981 . Population dynamics of wild
brown trout and associated sport fisheries in four Central
Wisconsin streams . Wis . Dep. Nat. Resour . Tech . Bull .
121 : 1-26 .
Donald, D . B . & D . J . Alger, 1986 . Dynamics of unexploited
and lightly exploited populations of rainbow trout (Salmo
gairdneri) from coastal, montane, and subalpine lakes in
western Canada. Can . J . Fish . aquat. Sci . 43 : 1733-1741 .
Donald, D . B . & D . J . Alger, 1989 . Evaluation of exploitation
as a means of improving growth in a stunted population of
brook trout . North Am . J . Fish . Mgmt. 9 : 177-183 .
Fraser, D . F ., D . A . DiMattia & J . D . Duncan, 1987 . Living
among predators : the response of a stream minnow to the
hazard of predation . In W . J . Matthews & D . C. Heins
(eds), Community and Evolutionary Ecology of North
American Stream Fishes . Oklahoma University Press,
Norman & London : 121-127 .
Garcia, A . & F . Braia, 1988 . Reproductive biology of brown
trout (Salmo trutta L .) in the Aller river (Asturias ; North-
ern Spain) . Pol . Arch . Hydrobiol . 35 : 361-373 .
Healey, M . C ., 1978 . Fecundity changes in exploited popula-
tions of lake whitefish (Coregonus clupeaformis) and lake
trout (Salvelinus namaycush). J . Fish . Res . Bd Can . 35 :
945-950 .
Healey, M . C ., 1980 . Growth and recruitment in experimen-
tally exploited lake whitefish (Coregonus clupeaformis) popu-
lations . Can . J . Fish. aquat . Sci . 37 : 255-267 .
Huntingford, F. A., N . B . Metcalfe & J . E . Thorpe, 1988 .
Feeding motivation and response to predation risk in At-
lantic salmon parr adopting different life history strategies .
J . Fish Biol . 32 : 777-782 .
Johnson, L ., 1976. Ecology of arctic populations of lake trout,
Salvelinus namaycush, lake whitefish, Coregonus
clupeaformis, arctic charr, S . alpinus, and associated species
in unexploited lakes of the Cannadian Northwest Territo-
ries . J . Fish . Res . Bd Can, 33 : 2459-2488 .
Jonsson, B ., 1977 . Demographic strategy in a brown trout
population in Western Norway . Zool . Scr . 6 : 255-263 .
Jonsson, B . & T . Sandlund, 1979 . Environmental factors and
life history of isolated river stocks of brown trout (Salmo
trutta m . fario) in Sore Osa river system, Norway . Envir .
Biol . Fishes 4 : 43-54 .
Lopez-Alvarez, J . V., 1984 . Observaciones sobre la aliment-
acion natural de la Trucha Comun (Salmo trutta fario) en
algunos rios de la cuenca del Duero . Limnetica 1 : 247-255 .
Mortensen, E ., 1977 . Population, survival, growth and pro-
duction of Salmo trutta in a small Danish stream . Oikos 28 :
9-15 .
McDonald, M . E. & A . E . Hershey, 1989. Size structure of a
lake trout (Salvelinus namaycush) population in an arctic
lake : influence of angling and implications for fish commu-
nity structure. Can . J . Fish . aquat . Sci . 46 : 2153-2156 .
Pielou, E . C ., 1969 . An introduction to mathematical ecology .
Wiley, New York.
Ricker, W . E ., 1950 . Cycle dominance among the Fraser
sockeye . Ecology 31 : 6-26 .
Scarnecchia, D . L ., 1983 . Trout and char production in
Colorado's small streams . Ph . D . Thesis, Colorado State
University, Fort Collins, Colorado .
Shetter, D . S ., 1969 . The effects of certain angling regulation
on stream trout populations . In T. G . Northcote (ed .),
Symposium on Salmon and Trout in Streams . University
of British Columbia, Vancouver : 333-353 .
Skurdal, J ., O . Hegge & T . Hesthagen, 1989 . Exploitation
rate, survival and movements of brown trout (Salmo
trutta L.) stocked at takeable size in the regulated rivers
Lagen and Otta, Southern Norway . Regulated Rivers : Re-
search & Management 3 : 247-253 .
Suarez, J . L., L. Reiriz & R . Anadon, 1988 . Feeding relation-
ships between two salmonid species and the benthic com-
munity . Pol . Arch . Hydrobiol . 35 : 341-359 .
Swales, S . & J . D . Fish, 1986 . Angling catch returns as indi-
cators of the status of upland trout lakes . Aquacult. Fish .
Mgmt 17 : 75-93 .
Werner, E . E ., J . F . Gilliam, D . J . Hall & G . G . Mittelbach,
1983 . An experimental test of the effects of predation risk
on habitat use in fish . Ecology 64 : 1540-1548 .
Zippin, C., 1956. An evaluation of the removal method of
estimating animal populations . Biometrics 12 : 163-189.