Fish Management in A Variable Water Envi

Fish management
in a variable water
environment
Fish management
in a variable water
environment
edited by:
Małgorzata Jankun
Grażyna Furgała-Selezniow
Małgorzata Woźniak
Anna M. Wiśniewska
Olsztyn 2011
Reviewers:
prof. dr hab. Jan Glogowski; prof. dr hab. Małgorzata Jankun-Woźnicka;
prof. dr hab. Janusz Terlecki; prof. dr hab. Teresa Własow; prof. dr hab. Zdzisław Zakęś
dr hab. Krystyna Demska-Zakęś, prof. UWM; dr hab. Roman Kujawa, prof. UWM;
dr hab. Andrzej Skrzypczak, dr hab. Małgorzata Woźniak, prof. UWM; dr Anna M. Wiśniewska;
dr Grażyna Furgała-Selezniow, dr Piotr Hliwa; dr Jarosław Król; dr Krzysztof Kupren
Translating: mgr Jolanta Idźkowska
The publication is co-financed under the EEA Financial Mechanism and Nor-
way Grants, priority sector „Environment and sustainable development”, the
Project no PL0468 „Ichthyological biodiversity of lakes - elaborating a model
for problem solution: a case study of natural resources of autochtonous com-
mon whitefish in Lebsko Lake (Lebsko Lake whitefish) (acronim Fish-WILL).
ISBN 978-83-60425-65-7
© 2011 Faculty of Environmental Protection and Fisheries

University of Warmia and Mazury in Olsztyn, Poland
Publishing: Agencja Wydawnicza „Argi” SC R.Błaszak, P. Pacholec, J. Prorok
Table of contents
Mamcarz A., Skrzypczak A. Between the ichthyological biodiversity
in the water bodies in Masuria (Mazury) and their management
– theories, problems and hopes. .............................................................................. 7
Szymańczak R., Sell J. Genetic monitoring of the fish stocking using
microsatellite DNA markers....................................................................................... 35
Zakęś Z., Demska-Zakęś K. Hatchery practice in the context of biodiversity
of aquatic ecosystems. ............................................................................................. 45
Sadowski J., Kempter J. Fish stocking versus biodiversity:
peaceful co-existence or continual conflict? . ........................................................... 53
Mickiewicz M., Wołos A. Species, type, and value of stocking material
released into Polish lakes from 2001 to 2009. . ........................................................ 65
Szmyt M. European grayling (Thymallus thymallus L.) stocking material
production in the context of the conservation of natural biodiversity
of the environment and active protection of the species. ........................................ 77
Turkowski K. Utility and market value of lakes. . ....................................................... 87
Skrzypczak A., Mamcarz A., Gierej A. Fishery classification of lakes
in the context of lake eutrophication and changes in the structure
of their ichthyofauna. .............................................................................................. 105
Szczepkowski M., Stabiński R. The state and prospects of the fish management
of coregonids in the context of changes in the fishery industry. .............................. 121
Martyniak A., Mierzejewska K., Stańczak K., Osewski M., Kisielewski K., Juńczyk M.,
Dziemian Ł., Zawal P. Effect of northern pike Esox lucius L. on a spawning
population of vendace Coregonus albula (L.) in Wigry Lake. . .................................. 127
Augustyn L. Formation of the ichthyofauna in the Czorsztyński Dammed Reservoir
(the Dunajec River system – southern Poland). ......................................................... 137
Tański A., Czerniejewski P., Keszka S., Pender R. Structure of ichthyofauna
in different fishery types of lakes used for angling by the Polish Angling Association,
the Region of Szczecin. ............................................................................................ 151
Wziątek B., Tandyrak R., Bowszys M. Biomanipulation as a method of creating
habitats for endangered fish species: a case study of the introduction of whitefish
into Lake Starodworskie. .......................................................................................... 157
Półgęsek M., Hofsoe P., Mysłowski B. Irresponsible aquarists
– a threat to native ichthyofauna? ........................................................................... 163
Marszelewski W., Solarczyk A. Changes in the oxygen status of lakes
and their effect on pelagic ichthyofauna. ................................................................ 167
Kubiak J., Machula S. Habitat conditions for the ichthyofauna
in the largest lakes in the West Pomerania Lake District. .......................................... 179
Nędzarek A., Tórz A., Kubiak J., Machula S. Habitat conditions for
the ichthyofauna in anthropogenic Szmaragdowe Lake (Szczecin, Poland)
in 2009-2010. .......................................................................................................... 193
Kujawa R., Pol P., Mamcarz A., Furgała-Selezniow G. Preliminary studies
on fishing and transport of the sichel (Pelecus cultratus L., 1758) for
reproduction under controlled conditions. . ............................................................. 201
Dziewulska K., Domagała J. Preliminary characterization of semen
of the salmon (Salmo salar L.) from rivers in West Pomerania. ................................. 207
Cejko B. I., Kowalski R. K., Kucharczyk D., Żarski D., Targońska K., Glogowski J.
Hormonal stimulation of males and its effectiveness in reproduction
and propagation of local populations of rheophilic cyprinid fish. ............................ 213
Sarosiek B., Kowalski R. Effect of cryoprotectants on activity of some enzymes
and motility parameters of spermatozoa of rainbow trout (Oncorhynchus mykiss)
and Siberian sturgeon (Acipenser baerii). ................................................................ 227
Żarski D., Targońska K., Krejszeff S., Palińska K., Piszczała S., Kucharczyk D. Effective-
ness of Ovaprim in controlled reproduction of some cyprinid fish species.
Part I: latency time and synchronization of ovulation. . ............................................ 237
Palińska K., Żarski D., Krejszeff S., Targońska K., Piszczała S., Kucharczyk D. Effective-
ness of Ovaprim in controlled reproduction of some cyprinid fish species.
Part II: gametes quality. . .......................................................................................... 245
Brzuska E. Differences in the efficiency of controlled reproduction of carp
Cyprinus carpio L. of the German line N, Hungarian line 7 and Hungarian line 8. ....... 251
Krejszeff S., Kucharczyk D., Targońska K., Żarski D., Kupren K., Mamcarz A.,
Piszczała S. Optimization of the reproduction of chub (Leuciscus cephalus L.)
under controlled conditions. .................................................................................... 265
Kirczuk L., Domagała J. Hybridization among fish and its importance to biodiversity. . .. 273
Between the ichthyological
biodiversity in the water
bodies in Masuria (Mazury) and
their management – theories,
problems and hopes
Andrzej Mamcarz, Andrzej Skrzypczak
Department of Lake and River Fisheries,

University of Warmia and Mazury in Olsztyn,
e-mail: mamcarz@uwm.edu.pl
Abstract
This paper contains a general discussion on ichthyological biodiversity, methods for
its evaluation and factors which can depress it on a global scale: transformations of the
natural environment and water contamination, introductions of alien species, overfish-
ing and global changes of climate. The biodiversity of the ichthyofauna in freshwater
bodies in the Masurian Lake District has been analyzed in detail: from the original state
of the post-glacial ichthyofauna through its transformations caused by ecological suc-
cession of lakes to the fishing exploitation of lakes, the course of introduction of new
species and fish stock releases. The authors pointed to the challenges facing institu-
tions which undertake nature conservation programmes concerning the ichthyofauna of
freshwater bodies in Masuria (climatic changes, legal issues regarding the management
of lands surrounding lakes and rivers, some negative aspects of the tourism, fishing and
angling use of waters and some adverse effects of aquaculture on decreasing biodiver-
sity of the ichthyofauna and stability of water ecosystems. It has been emphasized that
the general public should be more broadly educated about the need to conserve and
protect the water environment.
Biodiversity of lake ichthyofauna – different scales of the problem

According to the definition of the UNEP (1992), biological diversity of biodiversity is
“the variability among living organisms from all sources, including, terrestrial, marine,
and other aquatic ecosystems, and the ecological complexes of which they are part: this
includes diversity within species, between species and of ecosystems”. With this defini-
tion taken as the basis, biological diversity is typically discussed on three different levels:

Fish management in a variable water environment
genetic, species and ecosystems. Thus, according to Noss (1990), biodiversity has three el-
ementary components: composition (different genes, species, biological and physiological
functions, habitats, assemblages and landscapes as basic units of biodiversity), structure
(patterns of biodiversity arrangements are established by the ways in which components
between different borders dividing cells, organisms, populations and habitats interact)
and functions (transfer of genes between species and in speciation processes, behaviour
of organisms and interactions between and within populations, the role of biodiversity in
shaping the biological productivity). Biodiversity is an essential element of the stability of
ecosystems (Ehrlich and Wilson 1991). As the functional relationships between species in
ecosystems are rather poorly recognized, in many studies the determination of biodiversity
is reduced to making an inventory, which just brings the problem closer to us but does not
reflect its essence. Some estimates indicate that one species is represented by around 220
populations (Hughes et al. 1997), which altogether yields from 1.1 to 6.6 thousand billion
populations on the Earth. At the same time, it has been assessed (taking into consideration
the disappearance of populations as a linear function of habitat loss) that in the rainforests
alone 1,800 populations are destroyed each hour (annually, 16 million populations). Evalu-
ation of biodiversity on a global scale is difficult because of the incompleteness of data
on different environments. The available data, however, suggest that globally the percent-
age of endangered terrestrial vertebrates ranges from 11 to 25%, whereas in the case of
aquatic species it varies from 13 to 65% (Table 1). This information makes it evident that
aquatic environments are more severely threatened that terrestrial ones.
Table 1. Proportion of terrestrial and water vertebrates globally threatened (after McAllister et al. 2001)
Group Proportion threatened (%)

Mammals – all 25
Land birds 11
Waterfowl (freshwater) 13
Turtles, tortoises and terrapins 38
Crocodiles 43
Amphibians 25 (estimated)
Freshwater fishes 33 (estimated)
Freshwater mammals (freshwater dolphins and otters) 65
Like in other environments, biodiversity in systems of inland waters is a reflection of
the interaction between the existing assemblages of species and environmental condi-
tions. Although rivers, lakes and wetlands contain just 0.01% of water on the Earth, they
make a disproportionately high contribution to the concentration of biodiversity (Abell
et al. 2002) (Table 2). At the same time, water bodies are at a much higher risk of losing
biodiversity than other environments. Fish alone are not only the most diverse group of
vertebrates, but they also make up the largest share of threatened species (Bruton 1995,
Leidy and Moyle 1998, Duncan and Lockwood 2001). Among 737 globally threatened
fish species, 83 (11%) live in Europe (Figure 1).
Table 2. Species richness of the world’s major environments (after McAllister et al. 2001)
Environment Area of world surface (%) No. Living species (%) Richness: %species/%area
Freshwater 0.8 2.4 3.0
Terrestrial 28.4 77.5 2.7
Marine 70.8 14.7 0.2
Symbiotic N.A. 5.3 N.A.

Between the ichthyological biodiversity in the water bodies in Masuria…
Between 1970 and 2002, the biodiversity of freshwater bodies in the world decreased
by 55%, whereas the biodiversity of terrestrial and marine ecosystems fell by 32% (Mil-
lennium Ecosystem Assessment 2005). Small water bodies and ponds, which are a ref-
uge to many rare invertebrates and vertebrates, are disappearing at a particularly rapid
rate. According to Oertli et al. (2005), the rate of their disappearance is between 50 and
90%. Assemblages of species (populations) are a final product of an initial pool of genes
and a variety of long-term evolutionary processes such as speciation, disappearance
and migration from other ecosystems. Fish
constitute nearly half of the total number
of vertebrates on the Earth. Higher diversity
is found in tropical lakes than in lakes situ-
ated in the moderate climate zone, where
a strong influence of glaciation on the colo-
nization of waters and speciation of species
is observable (Amarasinghe and Welcomme
2002). Changes in aquatic environments af-
fect distribution of ichthyofauna as well as
a direction of its evolution.
Among numerous reasons why biological
diversity in water bodies declines, an essen-
tial one is a change in a habitat caused by Figure 1. Globally threatened fish species by region
humans, due to land use, urbanization, infra- (after GEO 3, 2002, recompiled) (N=737).
structure development and industrialization.
The present-day fragmentation of the landscape in which water bodies are found has
no precedence in history (Dale 1997). Aquatic systems are increasingly more often iso-
lated and separated. Many species living in specific habitats disappear rapidly due to such
changes. In transformed water ecosystems, between 20 to 35% freshwater fish are vulner-
able to extinction, threatened with extinction or becoming extinct (Braga et al. 1998).
Another considerable cause of loss of biodiversity is introduction of foreign or invasive
species, which can successfully compete with indigenous species and as a result cause
their disappearance (Mack et al. 2000, Lodge 2001, Blanchet et al. 2010). Alien species
can appear in waters because of incidental or thoughtless introductions. Development of
aquaculture and import of many new species to be cultured under controlled conditions
favour such introductions, as aquaculture facilities merge with the natural environment
(Fleming 1995). Braga et al. (1998) emphasize that aquatic ecosystems are especially vul-
nerable to such effects as a result of transformations of their environment and surround-
ings. In some cases, the same species is considered as being threatened in one location
and as an invasive one in another (Kozhara et al. 2007). Welcomme (1988) reports that
168 fish species representing 37 families have been introduced in different parts of the
world, of which at least 67 colonized new waters and 27 species have become an undesir-
able component in the environment. An increase in the number of introduced species was
particularly rapid in the second half of the 20th century (Figure 2).
Experts emphasize that species transfer between ecosystems within water bodies in
one country should be treated with equal care as international transfer (Kottelat and Whit-
ten 1996). The best-known case of catastrophic changes in a lake environment caused by
introduction of a new species occurred Lake Victoria in Africa (Stiassny 1996, Balirwa et

al. 2003). In the 1970s there

were over 300 endemic
species of Cichlidae, repre-
senting 99% of the lake’s
ichthyofauna. When the
Nile perch (Lates niloticus)
had been introduced to the
lake, most of the endemic
cichlids became extinct and
today they make up just 1%
of the ichthyofauna.
Likewise, water contam-
ination leads to a rapid loss
of biological diversity. Pol-
lutants disrupt the primary Figure2. Cumulative index of aquatic introductions (after GEO 3, 2002).
productivity of ecosystems;
they have an impact on the availability of nutrients and environmental processes by
modifying life conditions and composition of bodies of various organisms (Perrings et al.
1995). Many contaminants create synergic toxic combinations which produce a persist-
ent effect and gradually reduce productive capacity of ecosystems, without demonstrat-
ing evident and rapid signs of danger.
Another reason why biodiversity is diminishing is the widespread and uncontrollable
way in which natural water resources are used, which leads to their overfishing (Regier and
Henderson 1973, Kahn 1997). Overfishing most often occurs when there is no owner of
the resources (seas and oceans) or when the owner (the state) fails to fulfill appropriately
its role as a resources guardian (inland waters). Overfishing may also occur when the rate
of exploitation of resources is faster than the rate of their natural recreation. Overfishing
causes changes in aquatic ecosystems, a development which is completely neglected be-
cause of the prevailing view of their function as a commercial resource. Due to mass fish-
ing of the so-called valuable or select species, their place is taken over by species which are
smaller or grow faster, which creates an illusion of sustainable productivity, but coincides
with deep restructuring of the trophic relations in a given ecosystem. In the long run, this
situation most frequently leads to a complete collapse of exploited fish shoals.
An important factor which globally influences all the known causes of decline in the
ichthyofauna biodiversity is climate change (Reid 2006). Scientists estimate that an aver-
age temperature on the surface of our globe will have increased from 1.5 to 6.8oC by 2100
(Mann et al. 1999, Houghton et al. 2001) (Figure 3). The result will be the disappearance
of cold-water fish from most of the area they cover today and expansion of warm-water
fish as the waters grow warmer. It is estimated that in American waters, if the temperature
rises by a few degrees, at least 20 fish species will disappear (Matthews and Zimmermann
1990, Covich et al. 1997, Jackson and Mandrak 2002). The latest estimates indicate that
the availability of water in many rivers across the world will largely decrease by 2070 due
to climatic changes and increased uptake by humans. It is predicted that over 10% of the
river ichthyofauna will vanish as a result of drastic transformations in the environment of
these rivers (Xenopoulos et al. 2005).
10
Various changes in the

aquatic environment have
driven nearly 20% of the
world’s freshwater ich-
thyofauna on the brink
of extinction (Moyle and
Leidy 1992). Some authors
even claim that this is an
underestimate (Bräutigam
1999). The IUNC Red List of
Threatened Species of 1996
classifies as being threat-
ened 734 fish species, of
which 84% are freshwa-
ter ones (Mc Allister et al.
1997). In Poland, there are
37 threatened taxa of lam-
prey and fish (Witkowski et
Figure 3. The reconstructed and measured average temperature in the
al. 2009). Many, especially northern hemisphere (after Mann et al. 1999).
anadromous species, are
classified as critically endangered or endangered. There are just 19 taxa, i.e. 29.7% of the
native ichthyofauna, which are not threatened. Between 1999 and 2009, the number
of critically endangered species practically remained unchanged (57%). To make things
worse, a worrying tendency has been noticed for the number of species in the other
categories of threatened animals to increase.
Measuring biodiversity is one of the most important tasks in ecology because of the
role it plays in selecting a strategy for environmental conservation. This is a very complex
issue, both in terms of choosing the right methods and measures and because of the
lack of long-term studies that could serve as a starting point for the currently undertaken
screening procedures. The complex nature of this question has been thoroughly present-
ed in the paper published by Bredemeier et al. (2007).These very same problems make it
extremely difficult to include the question of biodiversity into the fishery practice (Longo
et al. 2007). The basic difficulty stems from the fact that there is no simple definition
or a way of measuring ichthyological biodiversity. The simplest measure of biodiversity,
applied in ecology, is the determination of the number of species in a habitat or a com-
munity. To this aim, the oldest biodiversity measure, such as the classical species diversity
index (H’) is applied. It is defined as:
H’ = - ∑ni /N · log2ni/N,
where: ni is the abundance of individuals belonging to the same species and N is the
total number of individuals (MacArthur 1955).
However, this measure is now considered to be overtly simplified, therefore indices
based on species richness and abundance are more frequently applied. Species richness
is the number of species in a community, while abundance explains the relative propor-
tion of each species in the environment. Species richness is measured with the diversity
index H (Shannon-Weaver Index) (Shannon-Weaver 1949):
11
H = - ∑pi ln pi,
where: pi = Ni/N (Ni = total number of individuals in species i), N = total number of
individuals of all species.
Species richness d (Margalef Index) is similar:
D = S – l/ln N,
where: S = total number of species, N = total number of individuals of all species.
Simpson’s Diversity Index (Simpson 1949) is used to measure abundance (dominance):
D = 1 – (∑pi 2),
where: pi is the proportion of the ith species in a sample.
The Non-Randomness Index E (Pielou’s Index) is another measure of biodiversity:
E = H/ln S,
where: H = index of diversity, S = total number of species.
The basic difference between the measures from the first and the second group is that
the species richness indices emphasize more strongly the role of rare species, whereas the
dominance measures – the role of the species which appear in higher numbers. Thus, the
choice of an index depends on what biodiversity components will be measured. If we wish
to pay attention to rare species, we should apply measures of species richness. However,
during the fishing exploitation, attention focuses on the most abundant species, therefore
dominance indices are applicable. Nonetheless, in order to apply Simpson’s index to fish-
eries, measure units need to be modified. Individuals as measures of a population must
be replaced by a share of biomass in total catches (Goda and Matsuoka 1986). Thus, the
ecological Simpson’s index can be expressed as (Gupta 2009):
2
Dt
s
Y it
i 1 Y t
where: Yit – is the catch of the ith species caught in time t, Yt – is the total catch in time
t and s is the number of species exploited in time t.
Application of this measure enables investigators to determine the relationship be-
tween the ichthyological diversity and total catches.
Conditions shaping the initial state of the ichthyological diversity in the water
bodies in Masuria
The subject of our analysis is the area of water bodies in north-eastern Poland, deline-
ated to the east and north by the state borders, to the west by the course of the Vistula
River and to the south – by the limits of the Baltic glaciation (Figure 4).
This area, like other post-glacial lands in north and central Europe, was a contact site for
the ichthyofauna originating from the basins of the Baltic, White and Caspian Seas. Many re-
searchers believe that the primordial neogenic ichthyofauna of Europe was very uniform and
consisted of such species as: sturgeon, perch, ruffe, burbot, two species of stickleback, salmon,
coregonids, grayling, roach, crucian carp, rudd, asp, bream, silver bream, bleak, dace, tench,
barbel, minnow, weather loach, bullhead, catfish, pike-perch, wild carp called sazan, ide,
12
spined loach, sunbleak. This ichthyofauna was characterized by the prevalence of cyprinids. As
a result of the glacial era, in the Quaternary, this group of fish was heavily diminished and then,
in the post-glacial era, complemented via migration of salmonids and representatives of the
Ponto-Caspian ichthyofauna. According to Alekseyev and Probatov (1969), following succes-
sive translocations of fish, the ultimate composition of the ichthyofauna in the areas bordering
with the region described hereby is represented by fish of five fauna complexes:
• Boreal highland complex (minnow, barbel, grayling, bullhead),
• Boreal lowland com-
plex (pike, crucian
carp, perch, ruffe,
roach),
• Former Tertiary com-
plex (catfish, wild
carp, weather loach)
• P o n t o - C a s p i a n
freshwater complex
(dace, blue bream,
rudd, asp, bleak,
barbel)
• Arctic freshwater
complex (European Figure 4. Masurian freshwaters.
whitefish, lake trout,
burbot).
With the existing hypotheses, archeological findings and later historical information tak-
en as a starting point, it can be assumed that during the post-glacial period the ichthyofauna
in the waters of Warmia and Masuria was represented by 50 species belonging to 15 families
(Table 3).
Nearly half of the spe-
cies in the original post-
glacial ichthyofauna con-
sisted of Cyprinidae, while
cold-water Salmonidae
made up a group of sub-
dominants (Figure 5). There
were 32 species classified
to 11 families inhabiting
the lakes (Figure 6).
As a result of a series
of processes, such as long-
Figure 5. Structure of the post-glacial ichthyofauna structure
term wasteful exploitation, in Mazurian freshwaters and the Vistula Lagoon (N = 50).
transformations and envi-
ronmental pollution or introduction of alien species and, possibly, too many needless
fish stock releases, at present 30 species are threatened, to a different degree, and one
species (Acipenser sturio) is considered extinct (Table 4). Lake trout, in turn, has disap-
peared from lakes in the Suwałki Lake District (Chybowski et al. 1994).
13
Table 3. Systematic account of original post-glacial ichthyofauna in the Mazurian freshwaters and Vistula Lagoon
(compiled from literature). Temperature preference after Blanck et al. 2007, supplemented (1 – stenotherm species
preferring cold waters, 2 – stenotherm species preferring cool waters, 3 – eurytherm species preferring cool waters)
Family Species Temperature preference

Petromyzon marinus (Linnaeus, 1758) 1
Lampetra fluviatilis (Linnaeus, 1758) 1
Petromyzontidae
Lampetra planeri (Bloch, 1784) 1
Eudontomyzon mariae Berg, 1931 1
Acipenseridae Acipenser sturio (Linnaeus, 1758) 1
Anguillidae Anguilla anguilla (Linnaeus, 1758)
Clupeidae Alosa fallax (Lacepede, 1803) 2
Alosa alosa (Linnaeus, 1758) 2
Barbus barbus (Linnaeus, 1758) 2
Carassius carassius (Linnaeus, 1758)
Gobio gobio (Linnaeus, 1758) 3
Tinca tinca (Linnaeus, 1758)
Rhodeus sericeus (Pallas, 1776) 3
Abramis brama (Linnaeus, 1758)
Abramis bjoerkna (Linnaeus, 1758) 3
Abramis ballerus (Linnaeus, 1758)
Vimba vimba (Linnaeus, 1758) 2
Rutilus rutilus (Linnaeus, 1758) 3
Cyprinidae Scardinius erythrophthalmus (Linnaeus, 1758)
Chondrostoma nasus (Linnaeus, 1758) 2
Aspius aspius (Linnaeus, 1758) 2
Leucaspius delinatus (Heckel, 1843)
Phoxinus phoxinus (Linnaeus, 1758) 2
Leuciscus leuciscus (Linnaeus, 1758) 2
Leuciscus idus (Linnaeus, 1758)
Leuciscus cephalus (Linnaeus, 1758) 3
Pelecus cultratus (Linnaeus, 1758) 2
Alburnus alburnus (Linnaeus, 1758) 3
Alburnoides bipunctatus (Bloch, 1782) 2
Cobitis taenia Linnaeus, 1758 2
Cobitidae
Misgurnus fossilis (Linnaeus, 1758)
Balitoridae Barbatula barbatula (Linnaeus, 1758) 3
Siluridae Silurus glanis Linnaeus, 1758
Esocidae Esox lucius Linnaeus, 1758 2
Osmeridae Osmerus eperlanus (Linnaeus, 1758) 1
Coregonus albula (Linnaeus, 1758) 1
Coregonus lavaretus (Linnaeus, 1758) 1
Thymallus thymallus (Linnaeus, 1758) 1
Salmonidae
Salmo salar Linnaeus, 1758 1
Salmo trutta trutta Linnaeus, 1758 1
Salmo trutta fario Linnaeus, 1758 1
Salmo trutta lacustris Linnaeus, 1758 1
Gadidae Lota lota (Linnaeus, 1758) 1
Pungitius pungitius (Linnaeus, 1758) 2
Gasterosteidae
Gasterosteus aculeatus Linnaeus, 1758 2
Cottus gobio Linnaeus, 1758 1
Cottidae
Cottus poecilopus Heckel, 1840 1
Perca fluviatilis Linnaeus, 1758 3
Percidae Gymnocephalus cernuus (Linnaeus, 1758) 3
Stizostedion lucioperca (Linnaeus, 1758) 3
14
Table 4. Taxonomic account of endangered and vulnerable ichthyofauna in Mazurian freshwaters and the Vistula
Lagoon (according to Witkowski et al. 2009). Threat criteria: EX – Extinct, EW – Extinct in the wild, CR – Critically Threat-
ened, EN – Endangered, VU – Vulnerable, NT – Near Threatened, CD – Conservation Dependent. Legal status: OG – pro-
tected in Poland, DS II – included in Annex II of Habitat Directive, DS V – included in Annex V of Habitat Directive.
Family Species Threat Legal status

Petromyzon marinus (Linnaeus, 1758) CR OG, DS II
Lampetra fluviatilis (Linnaeus, 1758) EN OG, DS II
Petromyzontidae
Lampetra planeri (Bloch, 1784) VU OG, DS II
Eudontomyzon mariae Berg, 1931 VU OG, DS II
Acipenseridae Acipenser oxyrhynchus (Mitchill, 1815) EX OG, DS II, DS V
Anguillidae Anguilla anguilla (Linnaeus, 1758) CD
Clupeidae Alosa fallax (Lacepede, 1803) CR OG, DS II, DS V
Alosa alosa (Linnaeus, 1758) CR OG, DS II, DS V
Barbus barbus (Linnaeus, 1758) VU DS V
Carassius carassius (Linnaeus, 1758) NT
Rhodeus sericeus (Pallas, 1776) VU OG, DS II
Vimba vimba (Linnaeus, 1758) CR/CD
Chondrostoma nasus (Linnaeus, 1758) EN
Cyprinidae
Aspius aspius (Linnaeus, 1758) NT DS II, DS V
Phoxinus phoxinus (Linnaeus, 1758) NT
Leuciscus leuciscus (Linnaeus, 1758) NT
Pelecus cultratus (Linnaeus, 1758) CR OG, DS II, DS V
Alburnoides bipunctatus (Bloch, 1782) EN OG
Cobitidae Cobitis taenia Linnaeus, 1758 OG
Misgurnus fossilis (Linnaeus, 1758) VU OG, DS II
Balitoridae Barbatula barbatula (Linnaeus, 1758) OG
Siluridae Silurus glanis Linnaeus, 1758 NT
Osmeridae Osmerus eperlanus (Linnaeus, 1758) VU
Coregonus albula (Linnaeus, 1758) VU DS V
Coregonus lavaretus (Linnaeus, 1758) VU DS V
Salmonidae Thymallus thymallus (Linnaeus, 1758) CD DS V
Salmo salar Linnaeus, 1758 EW/CD DS II, DS V
Salmo trutta trutta Linnaeus, 1758 CD
Salmo trutta fario Linnaeus, 1758 CD
Gadidae Lota lota (Linnaeus, 1758) VU
Cottidae Cottus gobio Linnaeus, 1758 VU OG, DS II
Cottus poecilopus Heckel, 1840 VU OG
Ecological succession
in lakes and its effect on
the biodiversity of ich-
thyofauna
After several glacia-
tions, the freshwater ich-
thyofauna of Europe is
evidently poorer than on
other continents (Lev-
eque et al. 2008). Since
the last glaciation, post- Figure 6. Structure of the post-glacial ichthyofauna
glacial lakes colonized by in the Mazurian Lakes (N=32).
15
fish have been undergoing long-term processes of ecological successions, vividly called
‘the aging of lakes’. Historical data clearly indicate that due to some changes in the bal-
ance between precipitations and water evaporation and runoffs, the location of lakes has
changed dramatically compared to the original state (Street and Grove 1979). Acceleration
of eutrophication and transformations of the ichthyofauna structure at an unprecedent-
ed rate caused by climatic changes has already been recorded in Scandinavian countries
(Degerman et al. 2001, Voutilainen and Huuskonen 2010). But the same process occurs in
Poland, where many lakes have disappeared. The data reported by Choiński (1991) proves
that between 1954 and 1991, 643 lakes in Masuria vanished and the total acreage of lakes
declined by 10%. The primary cause of their disappearance is increased eutrophication,
resulting mainly from more intensive farming on lands around lakes. The size structure of
Masurian lakes is exceptionally unfavourable, because 88.5% cover no more than 100 ha
(Figure 7). Considering the rate at which the lakes have been disappearing until now as
well as the predicted climatic changes over the next 100 years, it can be feared that a vast
majority of these lakes will vanish. Blanck et al. (2007) have made an inventory of environ-
mental preferences of 24 European freshwater fish species, which shows that as many as
40 out of 50 species which make up the autochthonous ichthyofauna of Masurian lakes
prefer cold and cool waters (Table 3). Thus, the disappearance of lakes will coincide with
the disappearance of 90% of species belonging to the existing post-glacial fauna, which
will be replaced by warm-water cyprinids and invasive fish.
The nature of succession of Polish lakes and their structure are of great significance to
institutions designing and implementing ecosystem monitoring and conservation strate-
gies (Mamcarz and Skrzypczak 2006). Because in most of the EU countries lakes less than
100 ha in surface area are considered to be too small and therefore unimportant for the
EU monitoring systems introduced under the Water Directive, the Polish administration
may feel in a position when they do not need to provide these lakes with constant care
but may wish to decrease the costs of monitoring open waters. In this situation, a large
proportion of ecosystems, inhabited by ichthyofauna, are not controlled by the owner
(the state) but left in the hands of leaseholders, who are business oriented.
However, an argument in favour of adapting a lake monitoring strategy is that during
ecological succession the primary ichthyological biodiversity is slowly being reduced and
consequently more valuable species with more stringent environmental requirements are
replaced by species of a broader ecological optimum. Such changes are known from the
now classical reports by Colby et al. (1972) and Hartman (1977). In respect of the Polish
lakes, there are corresponding changes in the structure of ichthyofauna in particular fish
type lakes (Figure 8). Tench-pike and crucian carp lakes, whose surface area is typically be-
low 100 ha, have very low species biodiversity, which encourages users to manage them
nearly the same way as ponds. But this leads to very rapid changes in the environment and
elimination of rare fish species, should there be any left, like Cobitidae, bitterling (Rhodeus
sericeus amarus) or other representatives of aquatic flora and fauna.
Effect of the management of open waters on biodiversity of ichthyofauna

Fishery exploitation
Archeological excavates dating back to the mid-Neolithic Age (4,000-2,500 B.C.) prove
that pike, perch and pike-perch were subjected to the strongest man-made pressure
(Makowiecki 2003, Antanaitis-Jacobs et al. 2009). Łęga’s (1949) analysis of the medieval
16
sources (12th-13th centuries)

pertaining to Pomerania,
a region neighboring with
the area discussed hereby,
suggests that the fresh-
water fishery in lakes and
rivers relied on pikes, stur-
geons and salmons. Eels
were also thought to be the
most attractive fish (Szc-
zygielski 1967). In Prussia,
the Teutonic Order held an
exclusive right for fishing,
either by its own fisher- Figure 7. Number of Masurian lakes in different
men or by issuing annual size categories (ha) (after Choinski 1991, modified).
licences (Olszewski 1948).
Fishing on open waters in
the Middle Ages involved
continual increase of the ex-
ploitation attained through
the improvement of fish-
ing techniques (mainly net
fishing). Rich and diverse
ichthyofauna encouraged
unlimited catches, prima-
rily during fish spawning
seasons. Thus, in the late
Middle Ages, the problem
of fish scarcity in rivers and
lakes began to occur, stimu-
Figure 8. The fish community structure in lakes according to the fishery
lating the mass develop- typology (after Zawisza 1965, modified). Lake types (1 – vendace-type,
ment of pond fish culture at 2 – bream-type, 3 – pikeperch-type, 4 – tench-pike type, 5 – crucian carp type).
the turn of the 15th and 16th
centuries (Szczygielski 1967). Pure exploitation of rivers and lakes continued until the 18th
century. The first inspections of the royal estates in the 1760s showed that lake fishery
gave a very small contribution to the proceeds from the rural economy. It turned into extra
jobs for peasants, who were granted a right to catch fish from feudal landlords in return
of a rent and later a lease rent. Fish poaching and thefts were very severely punished. The
same situation persisted after the partitions of Poland. Until World War One, no modern
techniques in inland open water fishery were practised in the Kingdom of Poland (Szczy-
gielski 1967). Consequently, the ichthyofauna of lakes and rivers was gradually devastated
due to the wasteful fishing exploitation and increasing pollution with industrial wastewa-
ter. In 1861, there were 950 lakes in the Kingdom of Poland and the production of river
and lake fish before World War One was just 900 tons annually (Szczygielski 1967). In the
former Polish lands partitioned by the Prussian Empire (in the Grand Duchy of Poznań), the
first fishery act, which was to halt the degradation of ichthyofauna in natural waters, was
17
issued in 1845. In the second half of the 19th century, lakes began to be stocked with fish
hatched in hatcheries. Since 1908, lakes were intensively stocked with eel.
During the twenty years of the interwar period, Poland held one of the top positions
in Europe in respect of the surface area of lakes. These lakes, however, are now beyond
our interest in this paper. The volume of lake fish production was estimated around 7,000
tons annually. For example, in Polesie the largest pikes and catfish were typically captured
during the spawning season. Fishery as an organized branch of economy received a firm
base as late as in 1932, when it was regulated by an act binding until 1985. The act sub-
ordinated the fishing production to rules of planned fishery protection. New regulations
on the fishery organization, leases, fishing limitations and fish stock releases were defined.
According to the Act of 1932, water bodies remained a state property and were leased to
various economic subjects by tender bid for the best lease rent (Różycki 1938).
Catches from the lakes in Masuria, in former East Prussia, reached about 2,500 tons
annually (Dąbrowski 1947). The fishery economy was extremely extensive. The catches
were dominated by small fish and bream, while select fish (whitefish, pike-perch) were
captured in small amounts (up to 40%). Pike-perch lived in just a few lakes.
After World War Two, when the state borders were shifted westwards, Poland lost
a large share of its lakes in the north-eastern territories but gained rich resources in former
East Prussia and Pomerania. In order to organize fishery in the new water bodies, new
organizational structures were established (State Fish Farms) alongside new research and
teaching units. In 1959, research was commenced to prepare lake management plans (sur-
vey plans) for about 2,000 lakes, based on scientific recognition of conditions in particular
aquatic ecosystems and adaptation of appropriate fishery management (Mamcarz and
Skrzypczak 2002). Plans for turning lakes into managed fish resources had to comprise an
evaluation of the current state of ecosystems and an outline of predicted activities in the
aquatic environment (Backiel and Zawisza 1965). At the same time, in order to find out
to what extent the economic results achieved until then reflected the potential produc-
tivity of water bodies, their users (State Fish Farms) were obliged to carry out analysis of
exploitation (Leopold 1968). This system of inland fishery management was modern and
unique in the world. It gave enormous possibilities of running sustainable fishery in lakes,
based on a scientific evaluation of production capacities of the environment. As was then
claimed that “planned economy when properly managed enables us to obtain production
output of maximum quantity and quality at the minimum outlays, maintaining its conti-
nuity” (Szczerbowski 1981). An important element of the fishery economy then consisted
of increasingly more intensive fish stocking and introductions of new species, which were
essential for increasing the fishing effectiveness. On the other hand, the fishery economy
in lakes was affected by political decisions (the state’s political system), which enforced
continuous production increase that was to prove the inherent superiority of socialist
economy. As a result, in many water bodies catches exceeded the limits set by the fishery
theory, thus degrading shoals of fish and disrupting the biological balance in ecosystems.
Nonetheless, this type of economy was thought to be rational.
The first five post-war years brought about a drastic increase in catches from lakes in
the Masurian Lake District, abundant in fish and exploited to a very small degree during
the war (Figure 9).
However, these catches seemed wasteful, which forced the institutions managing
the fishery industry to impose exploitation limits (Dąbrowski and Jaworski 1967). After
18
a period of rapid decline, the continuous increase in the subsequent years (up to 1975)
was a result of the advance in fishing tools and more intensive fish releases. In the
early 1960s, new, extremely efficient steelon nets were introduced (Dembiński 1962).
The years 1960-1975 witnessed very broadly planned releases of the fish stock of eel,
coregonids, carp and herbivorous fish. However, since 1975, a constant decrease in fish
catches has been observed. A study completed by Zawisza (1973) reveals that as soon as
the early 1970s the exploitation of lakes in Poland was two-fold higher than the allow-
able level of average annual production.
After 1989, when the process of liquidating the state lake fishery started and lakes
began to be privatized, new lake users were legally obliged to keep fishery plans as
a basic document for lake resources exploitation. However, the Ordinance of the Minis-
ter for Agriculture and Food Economy of 10 December 1997 (Ordinance 1997) did not
define clearly if the fishery management of lakes should be based on scientific grounds,
stemming from the current analysis of the environment. It stated that rational fishery
economy should be carried out in accord with fishery plans, leaving the decision how
they should be prepared to lake users. In turn, tender bids for leases of open waters
are based on competitively declared levels of fish stock releases, which disagrees with
the basic knowledge on ecological balance of water systems and their environmental
capacity. The situation is extremely bad in rivers, into which the fish species attractive to
anglers are introduced (carp, predatory fish), paying no respect to the fragmentation of
habitats where endangered river fish species dwell.
Treating open waters as an area of widespread economic activity, without imposing on
their users some tasks related to conservation of species biodiversity (e.g. an obligation to
protect genetically pure spawning flocks, limits or fishing restrictions, determination of an
effective size of a population below which catches should be postponed) will lead to fur-
ther degradation of existing fish shoals. The fact that such a development is likely to occur
can be implied by the situation in which many species are protected (or close to extinction)
and, at the same time, exploited (Table 5). In respect of some fish (eel), we have reached
a situation where a ban on
catching this fish may have
to be imposed all over the
EU. Future existence of
this species depends on
fish stock releases, but the
fish stock can be obtained
only if we allow adult fish
(commercially most valu-
able ones) to return to
the sea to their spawning
grounds. It is most prob-
ably too late to re-create
a stable population of this
species, supplying our in- Figure 9. Fishery exploitation of the Masurian Lakes
land freshwaters. in the second half of the 20th century.
19
Table 5. Taxonomic account of both exploited and protected ichthyofauna

in Mazurian freshwaters and the Vistula Lagoon. Keys as in Table 4.
Family Species Threat Legal status

Anguillidae Anguilla anguilla (Linnaeus, 1758) CD
Cyprinidae Barbus barbus (Linnaeus, 1758) VU DS V
Carassius carassius (Linnaeus, 1758) NT
Vimba vimba (Linnaeus, 1758) CR/CD
Chondrostoma nasus (Linnaeus, 1758) EN
Aspius aspius (Linnaeus, 1758) NT DS II, DS V
Pelecus cultratus (Linnaeus, 1758) CR OG, DS II, DS V
Siluridae Silurus glanis Linnaeus, 1758 NT
Salmonidae Coregonus albula (Linnaeus, 1758) VU DS V
Coregonus lavaretus (Linnaeus, 1758) VU DS V
Thymallus thymallus (Linnaeus, 1758) CD DS V
Salmo salar Linnaeus, 1758 EW/CD DS II, DS V
Salmo trutta trutta Linnaeus, 1758 CD
Salmo trutta fario Linnaeus, 1758 CD
Gadidae Lota lota (Linnaeus, 1758) VU
Fish introductions
Carp was the first fish to be introduced in the Masurian Lake District. Łęga (1949) reports
that first ponds began to appear in Pomerania in the 13th century. The first mention of fish
ponds near Tczew dates back to 1258. The author lists 11 villages in East Pomerania, where
fish ponds were kept. Although he does not say whether they were used for carp (Cyprinus
carpio L.) culture, it can be assumed that that was the beginning of its introduction in north-
ern Poland. Further expansion of this species to the north and the east of the area settled by
Baltic Prussians was connected with the gradual conquest and colonization of this land by the
Teutonic Order. Quoting the German chronicler Hennenberg, Olszewski (1948) reports that
carp ponds existed in Prussia as early as 1398. What is more, he cites a surprising fact that carp
had been known and cultured in Masuria even before the Teutonic Knights arrived. Inventories
made for the Teutonic castle in Klaipeda prove that carp did not appear there until the 15th
century as food consumed by Teutonic monks (Žulkus and Daugnora 2009). In 1402, 28,800
carps were eaten at the castle (Willoweit 1969 after Žulkus and Daugnora 2009).
German papers (Seligo 1912 after Bernatowicz 1948) prove that in 1880 there were
no populations of autochthonous Baltic whitefish in any of the lakes in East Prussia. Be-
necke (1880) mentions artificial breeding and fish stock releases of Coregonus lavaretus
maraena and Coregonus fera to Masurian lakes. Lake leasers imported the fish stock
from different places (Bodensee, Miedwie, Salentersee, Peipus and the Curonian Lagoon)
as they wished. Most of the whitefish fish released was from Peipus Lake (Coregonus
lavaretus maraenoides Polyakov) in Russia because in 1905-1917 it was the sole fish
stock material (Bernatowicz 1948, Rolik and Rembiszewski 1987). Since 1921, the lakes
were stocked exclusively with whitefish from Gołdapiwo Lake (dense-filter form). Ac-
cording to Wiese (1937), before WW2, Baltic whitefish was present in 46 lakes. In 1937
and 1938 fish fry of whitefish from Peipus Lake was also released to Wigry Lake, at the
amount of 500 individuals/ha. In 1948-1949, 134 adult individuals were caught in this
lake. After the war, several lakes near Giżycko were stocked with this fish species.
Germans also introduced different forms of trout and their hybrids. As Benecke reports
(1880), since 1872 lake char (Salvelinus alpinus, according to former terminology Salmo
20
salvelinus) was produced for stocking open waters. In 1880, Max von dem Borne imported
rainbow trout (Oncorhynchus mykiss Walbaum) from the USA to Germany, and in 1881-
1889 introductions of this species began in Pomerania and East Prussia. The aquaculture
of rainbow trout in Poland started in 1904, when eyed spawn of this fish was imported
from Sweden to a hatchery in Złoty Potok (Sakowicz 1952). Following these initial intro-
ductions to rivers and escapes from ponds, rainbow trout reached many ecosystems of
natural waters (including the rivers Wadąg, Bauda, Pasłęka, Wałsza, Drwęca and Czarna
Hańcza). As Witkowski claims (1996), rainbow trout can be found in 25% of the Polish
rivers. Brook trout (Salvelinus fontinalis) was brought to Poland from Germany in 1890. It
can be found in water bodies in Masuria, for example in the Bauda River.
Golden carp (Carassius auratus) has been known in Europe since 1611 and has been
present in European waters since 1728 (Bolle 1886). This fish was very popular to keep in
palace gardens and fish aquaria, from which it most probably spread to open waters. It is
likely that it was brought and cultured in ponds at palaces of the aristocracy in former East
Prussia. In the 19th century, its presence was recorded in waters near Gdańsk (Rathke 1824)
and in Prussia (Lorek 1837). In Poland, it appeared in the mid 1930s, most probably brought
accidentally with fish stock (Gąsowska 1934). Later, it spread all over Poland and to the Mas-
urian Lake District. The expansion was encouraged by fish stock releases in some lakes.
Brown bullhead (Ictalurus nebulosus) was brought to eastern Poland (the farm Wielko-
ryta) in 1920-1925 (Sakowicz 1952), from where it gradually spread over an increasingly
larger area. The latest data (Grabowska et al. 2008) state that this species is present in the
south-northern part of the Masurian Lake District. It can be found in the Drwęca River, too.
In 1958, introduction of ripus (Coregonus albula ladogensis Pravdin) to Masurian
lakes (Maróz, Klawój, Sarż) started. In 1960, the fish were released to Radomno Lake
near Iława (Gottwald 1960). The result of the introduction is unknown. It is interesting
that at the same time when ripus was brought from Russia, its hybrid with Baltic white-
fish Coregonus albula ladogensis x Coregonus lavaretus) began to be introduced to the
Masurian Lakes. In 1958, eyed spawn of this hybrid was imported from Russia. After
incubation and rearing, fish fry weighing around 8 g was introduced to Dgał Wielki Lake
near Giżycko. After 3 years, natural spawning of the introduced fish was observed. No
further introductions were done (Bernatowicz 1964).
In 1969, an accidental introduction of European mudminnow (Umbra krameri Wal-
baum) to Kiernoz Lake was observed to have taken place when the lake was stocked
with carp and tench from Hungary (Ejsymont and Słoniewski 1969). In the following
years, this species has not been observed in the Masurian Lakes.
In the 1960s four other species were introduced to the Masurian Lakes. In 1964,
grass carp (Ctenopharyngodon idella) was introduced and a year later – two species of
silver carp (Hypophthalmichthys molitrix and Aristichthys nobilis) (Krzywosz 2000a, b).
In 1966, peled (Coregonus peled) was introduced to the lakes in Masuria. In 1966-1983,
fish introduction encompassed 181 lakes of different size (Mamcarz 1992a).
In the 1990s, several species of sturgeon were imported to Poland as new culture
fish. In closed fish rearing facilities the following sturgeons were kept: Siberian (Acipens-
er baeri), Russian (A. gueldenstaedti), stellate (A. stellatus) and a hybrid A. ruthenus x H.
huso, called bester (Kolman 1993, 2000). What is of interest here is that all these species
are endangered or critically endangered in their natural habitats (Birstein et al. 1997). In
21
Poland, they are cultured, inter alies, in the fish rearing farm in Dgał near Giżycko. Ever
since the sturgeon culture started, presence of single individuals of these fish have been
noticed in natural waters (Keszka and Stepanowska 1997). It is therefore highly probable
that they can appear in the region analyzed in this paper.
In 1984-1985, attempts were made to introduce to the Masurian Lakes another repre-
sentative of Siberian coregonids (Coregonus muksun). The fish were imported from Finland
and released to Dgał Wielki Lake, however the introduction was unsuccessful (Uryn 1987).
In the 1960s, it was also attempted to introduce endemic lake trout (Salmo trutta
lacustris) from Wdzydze lake to rivers in Masuria (Dąbrowski and Jaworski 1967). Lake
trout was released for instance to Pasłęka River, Pierzchalski Reservoir, the Wadąg River
and Wadąg Lake and the Czarna Hańcza River. However, no positive effects were ob-
tained (Grudniewski, unpublished data). In the 1990s, it was introduced to lakes in the
Suwałki Lake District (Chybowski et al. 1994).
Fish introductions running for many years have resulted in the appearance of 27 new
species in Masurian waters, of which some failed to acclimate and died out and some,
like coregonids, crossed with local forms so that their present status is unknown (Table 6).
Some of the new species (Acipenseridae) are cultured in closed aquaculture facilities and,
except Acipenser oxyrhynchus, their presence in open waters is not commonly verified.
Table 6. Taxonomic account of ichthyofauna introduced into Mazurian fresh waters.
Family Species Year (Period)

Salmonidae Oncorhynchus mykiss Walbaum, 1792 1881-1889
Salmo trutta lacustris Linnaeus, 1758 1962, 1991
Salvelinus alpinus Linnaeus, 1758 1872
Salvelinus fontinalis (Michill, 1815) After 1890
Coregonus lavaretus maraenoides Polyakov, 1874 After 1878
Coregonus lavaretus maraena Bloch, 1779 1872
Coregonus lavaretus holsatus Thienemann, 1916 After 1878
Coregonus lavaretus generosus Peters, 1875 After 1878
Coregonus wartmanni (Bloch 1784) After 1878
Coregonus fera Jurine 1825 1872
Coregonus peled Gmelin, 1788 1966
Coregonus muksun Pallas, 1776 1984
Coregonus albula ladogensis Pravdin, 1939 1958
Coregonus albula ladogensis x Coregonus lavaretus 1958
Cyprinidae Cyprinus carpio Linnaeus, 1758 After 1398
Carassius auratus (Linnaeus, 1758) ?
Carassius auratus gibelio (Bloch, 1783) About 1837
Ctenopharyngodon idella Valenciennes, 1844 1964
Hypophthalmichthys molitrix Valenciennes, 1844 1965
Aristichthys nobilis Richardson, 1845 1965
Umbridae Umbra krameri Walbaum, 1792 1969
Acipenseridae Acipenser baeri Brandt, 1869 1992
Acipenser gueldenstaedti Brandt, 1833 1992
Acipenser stellatus Pallas, 1771 1992
Acipenser ruthenus x Huso huso 1992
Acipenser oxyrhynchus Mitchill, 1815 2006
Ictaluridae Ictalurus nebulosus LeSueur, 1819 ?
22
Another effect of the planned and incidental introductions will be an obviously increas-
ing presence of invasive species, such as the gobies (Gobiidae), brown bullhead (Ictalurus
nebulosus) and topmouth gudgeon, as well as other fish from aquaculture facilities and
aquaria. Among the species which can become endangered in the nearest future all over Po-
land is sunbleak (Leucaspius delineatus). As a consequence of the rapid expansion of the in-
cidentally introduced topmouth gudgeon (Pseudorasbora parva) observed now in Poland, it
can be expected that sunbleak will become completely extinct. The cause is a fatally danger-
ous parasite carried by this invasive species. The new parasite will probably be a new threat
to the ichthyological diversity of European waters and aquaculture (Gozlan et al. 2005).
Some authors (Padhi and Mandal 1994) point to another threat to ichthyological biodi-
versity, namely wrong hatchery practice, which leads to the crossing of species and produc-
tion of hybrids, which are then released or spread to the natural environment. In Warmia
and Masuria (Warmia and Mazury), such a situation was widespread, first in respect of Bal-
tic whitefish (in the 19th c. and the first half of the 20th c., German fish releases with differ-
ent forms of whitefish) and later – the introduced peled (Coregonus peled Gmel.), which
– when being artificially reproduced – was crossed on a mass scale with Baltic whitefish
(Coregonus lavaretus L.)
and then settled in natural
waters (Mamcarz 1992b,
Luczynski et al. 1999).
At present, it can be
assumed that the ichthyo-
fauna of open waters in
Masuria (including the in-
troduced fish) consists of
60 species, half of which
are endangered to a vari-
ous extent (Figure 10). Figure 10. The current ichthyofauna structure of Masurian freshwaters.
Fish stock releases

Over the past two centuries, fish stock releases to lakes have been perceived as a basic
method for improving the efficiency of commercial fish catches or attractiveness of lakes as
game fish resources. Thus, the decision which water bodies to stock has never been based
on an evaluation of the ecological balance of ecosystems but strongly motivated by the wish
to strengthen the species of the then highest economic value. Fish stock releases aiming to
maintain balance between fish populations or to support their occurrence in lakes did not
start until it was noticed that many species were rapidly vanishing (no natural spawning).
Before World War Two, lakes were intensively stocked with eel and tench (Dąbrowski
and Jaworski 1967). Likewise, the conservation of ichthyofauna was on a high level. Af-
ter the war, the first mass fish stock releases started in 1945-1946, by stocking 21 lakes
with eels, vendace, pike and pike-perch. In the subsequent years, the range of fish stock
releases grew very quickly. In 1964, 13 fish species were stocked and during the peak
exploitation period, lakes were stocked with 17 species; after 1989, 20 species have been
reared to stock lakes and rivers (Table 7).
23
Table 7. Taxonomic account of ichthyofauna stocked to Mazurian fresh waters

after World War Two (a - after Dąbrowski and Jaworski 1967, modified).
Family Species 1964 - a 1975 After 1989

Anguillidae Anguilla anguilla (Linnaeus, 1758) x x x
Cyprinidae Carassius carassius (Linnaeus, 1758) x x x
Carassius auratus gibelio (Bloch 1783) x
Tinca tinca (Linnaeus, 1758) x x x
Abramis brama (Linnaeus, 1758) x x x
Rutilus rutilus (Linnaeus, 1758) x x x
Cyprinus carpio Linnaeus, 1758 x x x
Ctenopharyngodon idella Valenciennes 1844 x x x
Hypophtalmichthys molitrix Valenciennes 1844 x x
Aristichthys nobilis Richardson 1845 x x
Leuciscus idus (Linnaeus, 1758) x
Aspius aspius (Linnaeus, 1758) x
Esocidae Esox lucius Linnaeus, 1758 x x
Salmonidae Coregonus albula (Linnaeus, 1758) x x x
Coregonus lavaretus (Linnaeus, 1758) x x
Coregonus peled Gmelin, 1788 x x x
Salmo trutta trutta Linnaeus, 1758 x
Salmo trutta fario Linnaeus, 1758 x x
Salmo trutta lacustris Linnaeus, 1758 x x
Oncorhynchus mykiss Walbaum 1792 x x
Percidae Stizostedion lucioperca (Linnaeus, 1758) x x
Perca fluviatilis Linnaeus, 1758 x x
Siluridae Silurus glanis Linnaeus, 1758 x
Gadidae Lota lota Linnaeus, 1758 x
Growing contribution of fish stock releases to open waters management has always
been perceived as a positive event, sustaining the economic effectiveness of running
business on open waters. Currently, however, another aspect of fish stocking has been
emphasized, namely its role in active protection of dying and endangered species (Lirski
et al. 2008). Paradoxically, some of these species, owing to their market value (salmo-
nids, eel) are nearly instantly caught from the lakes before they manage to shape an
effective structure of reproducing populations. Intensive fish stock releases alongside
intensive fishing exploitation are not neutral to the equilibrium and structure of ecosys-
tems, which are losing their stability. Thus, we can speak of “a paradox of protected spe-
cies in unprotected ecosystems”. Currently, 15 endangered species out of all 32 natural
representatives of the Masurian ichthyofauna are protected in this manner (Table 5).
American studies on lakes in Minnesota have implied an ambiguous role of fish stock
releases (Radomski and Goeman 1995). Fish releases have raised the species richness but
at the same time have led to the structural homogenization of many lakes. The same
must be true about our lakes, because decisions to stock them are first of all dictated by
economic considerations (valuable food fish) and not ecological one. Thus, it is difficult
to speak about the actual conservation of ichthyological biodiversity, which would be
based on an appropriate diversity of habitats.
Where we are heading – or reaching a compromise

Among the challenges we face on the way of improving the conservation of biodiver-
sity of waters, global climatic changes prevail. However, the question of total degradation
24
of biodiversity is more and more often analyzed in terms of economic transformations

and a choice of the way in which economic growth should be attained (Clausen and York
2008). The global character of this problem creates serious difficulties in reaching a com-
promise so as to undertake successful action limiting negative changes. In short, we may
sooner experience a global catastrophe than be able to reach a global understanding.
Likewise, the ambitious plans of the European Union with respect to improving the state
of natural water bodies in the member countries and halting the loss of biodiversity until
2010 cannot be accomplished without some radical improvement in the state of the river
and lake environment on a local level. Many EU legal initiatives concerning conservation
of the ichthyofauna biodiversity have not been fully implemented yet or supported by the
authorities of particular member states (Greenstreet 2008, Hiddink et al. 2008). As some
researchers suggest (Strauss and Pezold 2009, cited after Kostoski et al. 2010), if the cur-
rent trend of biodiversity loss continues, half of all the species will become extinct in less
than a hundred years. Until now, however, most of the activities undertaken in many coun-
tries to reverse this trend are limited to monitoring the resources and making small efforts
to reduce its negative effects. Active reconstruction of aquatic ecosystems is affordable to
the wealthiest countries only. As Evans (2009) reports, in 2006 the USA allocated 62.8 mil-
lion dollars for the conservation of two freshwater fish species (Scaphirhynchus albus and
Salvelinus confluentus) and 320 million dollars to protect two migrant species of salmon
(Oncorhynchus tshawytscha and Oncorhynchus mykiss). The list of various threats to the
biodiversity of freshwater aquatic ecosystems is long (Table 8).
Some can be limited by establishing Freshwater Conservation Areas (Suski and Cooke
2006). Some researchers (Olden et al. 2010) point to the need of improving the applied
nature conservation strategies, which far too often refer to systems of protecting terres-
trial organisms and therefore are not very successful in respect of the water environment.
The role of continuity of habitats (conservation area networks) as a basis for effective con-
servation of ecological processes in water ecosystems is raised (Hermoso et al. 2010). It is
also emphasized that we need to learn more about ecological theories on populations and
include them into decision making processes regarding planned ichthyofauna conserva-
tion. Another important factor which makes it impossible to prepare a proper conserva-
tion strategy is the lack of knowledge on the minimum size of populations an d minimum
habitat requirements for most of the freshwater fish (Nel et al. 2009).
A great challenge to the conservation of biological diversity of rivers and lakes is the
proper conservation and/or reconstruction of their shore zones and the land management
around water bodies that is in accord with the principles of nature conservation (Lake et al.
2000). In this area, however, principles of ecology are very quickly outcompeted by political
and economic objectives. Despite the commonly expressed approval for eco-friendly activi-
ties concerning water bodies, in practice the regulations are circumvented, local aims are
preferred to higher values and river or lake shores as well as surrounding land are wastefully
“utilized” by various economic entities. Such actions are very often motivated by promotion
of tourism as a way of stimulating the economic activity in a given region. This is dangerous
because the development of aquatic tourism is not supported by analyses of tourist load
on the environment, which could help to make rational decisions on localization of tourist
facilities and regions of intensive tourist concentration. Negative experience in promoting
uncontrollable tourism development, known from other European countries (Kostoski et al.
2010) should not be duplicated in the Masurian Lake District, where, in many places, the
25
tourism industry has started to develop yet. Angling, which also has a global scale, is one of
the basic components of aquatic tourism and recreation. At present, in 22 European coun-
tries there are at least 21.3 million anglers, who spend large amounts of money every year
on their sport (Cowx 2002). The pursuit of their hobby brings a considerable contribution to
the economies of many countries, including the fishery business, which in many parts of Eu-
rope is in decline. Angling helps to develop fish hatchery, to stimulate broad-scale fish stock
releases to open waters and to evidently replace the fishing use of waters by angling. This
development, by many decision-makers thought to be a very positive trend, animating the
regional economy, has a very negative influence on the natural environment and ichthyo-
logical biodiversity (Cambray 2003). Angling plays a particularly strong and disgraceful role
in global spreading of invasive species, destroying the local ichthyofauna (Table 9).
Table 8. Anthropogenic and environmental threats for freshwater ecosystems reducing biodiversity and ecosys-
tem integrity (after Suski i Cooke 2006, modified). Ranking of severity threat (L – low, M – moderate, H – high),
amount of protection offered by Freshwater Protected Areas (FPAs) (L – low, M – moderate, H – high).
Severity Protection
Threat Comments
of threat from FPA
Species introductions H L Introduced species may outcompete endemics
Dams/weirs/barriers H M Prevent movements and isolate populations
Can change water temperatures to exceed
Global warming H L
thermal maximum of species
Shoreline Can modify spawning/rearing areas and
H H
development change water temperatures
Excessive harvest can negatively influence
Commercial harvest H H
populations
Population declines due to recreational
Recreational fishing M H
angling (Post et al. 2002)
Excessive harvest can negatively influence
Artisanal fishing H H
populations
Differences in water levels can negatively
Flow regulation H L influence spawning/rearing areas, altered
behaviour and energetic costs
May help promote the popularity of closed
Tourism L H
areas
Changing nutrient levels can disrupt food
Eutrophication H L
webs
Acid rain has damaged many freshwater
Acidification H L
ecosystems
Increased UV radiation can negatively influ-
UV radiation M L
ence many littoral processes/organisms
Boating
L H Can be problem in shallow environments
sedimentation
Atmospheric Many potential pollutants can enter fresh-
M L
pollutant deposition water environment from atmosphere
Runoff and tailings can negatively influence
Mineral abstraction M L
freshwater environment
26
Table 9. Eight fish species listed in the “One Hundred of the World’s Worst Invasive Alien Species”
(after Cambray 2003).
Species Reason for introduction Impact

Aquaculture but also
Clarias batrachus Indigenous fish and other aquatic biota
for sport fishing
Aquaculture but also Reduces water quality and destroys
Cyprinus carpio
for sport fishing aquatic vegetation by uprooting it
Mosquito control
Gambusia affinis (indigenous species could Harmful due to predaceous habits
also have done this)
Contributed to extinction of 200 fish spe-
Lates niloticus Food and sport fishing cies in Lake Victoria resulting in devastat-
ing environmental impacts
Micropterus sal- Impact on indigenous fish, crayfish, am-
Sport fishing
moides phibians and insects
Displace indigenous species by competi-
Oncorhynchus
Sport fishing tion and predation, also impact on aquat-
mykiss
ic invertebrates
Oreochromis Aquaculture also Omnivorous eats almost anything from
mossambicus for sport fishing algae to insects
Severe impact on indigenous fish (espe-
cially other salmonids), amphibians, inver-
Salmo trutta Sport fishing
tebrates through predation, displacement
and food competition
The fishery economy should be based on a sustainable use of the ichthyological re-
sources, which will enable us to take advantage of long-term utilization of fish flocks and,
accordingly, provide long-term employment in this sector. Several elementary principles
must be obeyed so as not to cause degradation of fish resources: 1) the exploitation
rate must be lower than the natural growth rate of fish populations, 2) each population
should be kept above the minimum size which guarantees its reproduction (effective size),
3) a population must not lose its ecological function, 4) genetic variation, size, age and
gender structure of a population must not be degraded, 5) the flock behavior must be
maintained. Although we have quite extensive knowledge on how to act in order to avoid
overfishing, in practice unsustainable use is frequently implemented, causing disappear-
ance of fish flocks, biological disequilibrium and decline in biological diversity. Moreover,
the cascade effect of disappearance of fish species regulating trophic pyramids is com-
pletely ignored, which leads to a chain disappearance of subsequent trophic links and
a deep destalibilization of ecosystems (Dunne and Williams 2009). Considering the variety
of functions that lakes play in economy, back in the 1980s (Leopold et al. 1980) it was
postulated to utilize them according to one of the six functions (reservoirs of pure water,
nature reserves, retention reservoirs, non-fishing recreational reservoirs, angling reservoirs,
fishing reservoirs). This concept has not been applied and today all the lakes, except the
ones located in nature reserves, are used for both fishing and angling.
The dynamic growth of aquaculture, which we witness today, creates an impression
that it is possible to resign from the current use of the natural ichthyofauna resources by
multiplying fish production on controlled systems. However, aquaculture has its “dark
27
sides”, too, in respect to the aquatic environment. As the specification presented by Tis-
dell (2006) shows, the effect of aquaculture on the ichthyological biodiversity of waters
can be considerable (Table 10).
The author shows a series of negative events, such as uncontrollable escapes of cultured
fish, narrowing of the natural genetic variation, effect of health-sustaining manipulations and
transport of cultured fish, appropriation of natural water bodies for aquaculture facilities.
Some of these phenomena, such as production of genetically engineered organisms, should
be carefully considered, in terms of ecology and ethics, by scientists and decision-makers
(Myhr and Dalmo 2005). Tisdell (2006) draws our attention to the lack of rules governing
the selection of the fish genetic material. This situation encourages creation of selection pat-
terns which favour keeping ownership rights to created fish forms by different companies
and increasing their profits, but does not help to maintain biodiversity. At present, however,
the biggest “dark side” of aquaculture is the fact that it consumes enormous resources of
fish protein. For aquaculture to thrive, some 30% of the global fish catches are turned into
protein for feeding aquaculture fish. It is estimated that aquaculture of predatory fish used
15% of the world’s fish meal supply in 1995 (Martinez 1995).
It seems that the governing principle in relations between aquaculture and fishery
should be the rule of wise self-limitation, which means that as more intensive methods
of controlled production of particular fish species develop, more restrictive limitations
on their acquisition from the natural environment should be applied. This would lead
to a gradual transformation of natural resources into protected “genetic stores”, neces-
sary for running aquaculture. Aquaculture, even in its intensive form, largely relies on
supplementing its base with spawners or genetic material obtained from the wild. Its
future growth and sustenance are possible provided the natural resources are kept on
an unchanged level. It is worth considering today if that does not mean that in many
aquatic ecosystems fishing exploitation should not be completely limited in order to
keep suitable spawning fish flocks in ecological balance. Examples of methods for cal-
culation of volumes of allowable fish catches (effective size of a population) preventing
overfishing, are cited in research papers (Prusevich 1972, Hansen et al. 2006). This ap-
proach means a complete change in our view of the current fishery economy. It does
not, however, involve the economic loss to business entities running fishing exploitation
because the new tasks set under the national system of biodiversity conservation should
be compensated for by the state, the owner of the water bodies. Genetic monitoring of
the effective size of populations as a measure of genetic stability and inbreeding degree
should also play an important role (Hansen et al. 2006).
It is also of utmost importance to educate the society on the conservation of ichthyo-
logical biodiversity. Researchers and mass media, when dealing with issues of nature con-
servation, more readily discuss the question of protecting mammals and birds, which seem
closer to the interest of people, than aquatic fauna and fish in particular (Richter et al. 1997,
Ricciardi and Rasmussen 1999). The main reason is that lower vertebrates (fish) and aquatic
fauna feature relatively less often in nature protection research. Analogously, the popular
knowledge on functions and balance in water ecosystems among the general public is
poor and often combined with the lack of appropriate sensitivity to nature or negligence of
threats. In conflict situations, when different forms of human activity threatening water eco-
systems must be limited, arguments about the superiority of people’s needs (social needs)
over the existence of live nature are quoted. It is commonly believed that natural waters are
28
self-reproducing stores of goods, from which everybody can draw as needed. This approach
to the co-existence with nature is confirmed by the very weak capability of enforcing law by
the state nature conservation services (fishery guards), few and underinvested, who work
being aware of the small effectiveness of the tasks they perform.
Table 10. Aquaculture practices and their consequences for biodiversity loss (after Tisdell 2006).
Practice Consequences
Translocations of fish species or varietes of Loss of indigenous fish species and other
fish with their accidental or deliberate release wild species due to competition, habitat
to wild disturbance and so on. Examples include
translocation of carp, tilapia and trout
Release (accidental or deliberate) of improved May alter the genetic composition of the wild
varietes of fish or transgenic varieties to the stock if they are sufficiently fit for survival
wild (Myhr i Dalmo 2005) in the wild and the releases are sufficient in
number (Muir 2005)
Narrowing of the diversity of the genetic The genetic diversity of farmed fish stock is
stock in aquaculture due to human selection often much less than the wild stock for which
of species and their varietes (Hulata 2001) it is a substitute or replacement. Market
extension and globalization are strong forces
working in favour of reduced biodiversity of
farmed organisms
Appropriation of habitat and space of areas Wild species excluded or partly excluded from
used by wild species for aquaculture and aquaculture areas. Lose food sources, shelter
destruction or significant alteration of habitat and breeding areas
Exploitation of wild aquatic fish and materials Because of loss of food sources for wild fish
to provide food for aquaculture organisms and over harvesting of targeted species, loss
of biodiversity in the wild may occur
Use of chemicals and antibiotics in Possible loss of some such fauna with
aquaculture may adversely affect local negative impacts on the food chain and
aquatic microfauna and macrofauna potentially therefore, on higher order species
(Beardmore et al. 1997)
Intensive collection of seed for aquaculture May threaten wild stocks or alter the genetic
ranching composition of these
Movement of objects (biological and non- Accidental or incidental introduction of new
biological) over considerable distances for pathogens, parasites or pests generally to new
use in aquaculture areas with biodiversity loss possible
Considering the complex nature of the problems related to the biodiversity of natural
waters as well as the historical, economic and civilization background, it would be dif-
ficult to suggest any list of priorities that could help to halt this centuries-long process of
nature degradation. Point I of the Preamble to the EU Water Framework Directive states
that “Water is not a commercial product like any other but, rather, a heritage which
must be protected, defended and treated as such.” This opinion is a point of reference
for all activities undertaken in respect to live water ecosystems by anybody and every-
body, from states to individual persons. If we refer to this statement on each occasion,
while undertaking different actions, posing a threat to the safety of nature in water
ecosystems, this could be a true measure of our intentions.
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34
Genetic monitoring of the fish
stocking using microsatellite
DNA markers
Robert Szymańczak, Jerzy Sell
University of Gdańsk, Department of Genetics,

Kładki 24, 80-822 Gdańsk
sell@biotech.ug.edu.pl
Abstract
Conservation of genetic diversity of natural fish populations, which are ‘supported’
by releases of artificially reproduced material, frequently calls for solving a series of
problems. We are presenting some implications of fish stocking to the genetic structure
of populations and possibilities of applying microsatellite DNA as a genetic marker for
the assessment of the effects associated with fish stocking and in the conservation of
ichthyological diversity.
Effects of fish stocking on the genetic structure of a population

Fish stocking programmes are commonly applied in practice and their aims are to
protect and restore local populations which are at the risk of becoming extinct, and to
alleviate effects of reduction or loss of productivity of populations due to damage, modi-
fication or degradation of habitats or other functional components of an ecosystem.
Other objectives of fish stocking are intentional introduction of an exogenous population
to a new ecosystem; short-term fish breeding in order to attain maximum fish catches or
breeding fish in a closed system for commercial purposes (Epifanio et al. 2001).
Fish stocking, however, may have a considerable effect on the structure of populations
of a species. Natural populations of a given species may be completely different from one
another in both phenotypes and genotypes. Such differences could result from the influence
of the natural environment on a given phenotype, not having any genetic basis, or they may
be caused by the effect of genetic drift on a population’s gene pool, or else – which seems
extremely important – they can be direct reflection of the population being adapted to the
local environmental conditions (Dannewitz 2003). In the latter case, we deal with a dynamic
process, which contributes to the maintenance or increase in the frequency of occurrence of
traits within a population that enhance its chances for survival and successful reproduction.
Such traits, called adaptive ones, are genetically conditioned and their expression is cor-
35
related with modifications in survivability and fertility. These traits are subjected to natural
selection (Taylor 1991). Adaptation to local environmental conditions can be mirrored as
variability of morphological and meristic characteristics (Řstbye i in. 2005), changes in be-
haviour (Lahti et al. 2001), reproduction biology (Sakamoto et al. 1999), course of life cycle
(Hutchings and Jones 1998) or resistance to pathogens (Taylor 1991).
Introduction of individuals originating from other than local populations, therefore
lacking certain adaptive traits, might have very serious adverse consequences. In an ex-
treme case, it may lead to damaging the genetic structure, which has been created in the
course of adaptation of a local population, due to introgression. This in turn depresses
the degree of adaptability of a local population to life in a specific ecosystem. This situa-
tion also creates the risk that a local population, which was supposed to be reinforced by
stocking, may become extinct (Dannewitz 2003, McGinnity et al. 2003). Certainly, it is also
possible that introduction of brood stock will have only a slight effect due to insufficiently
low adaptability skills of introduced individuals. However, a study completed by McGinnity
et al. (2003) proved that introduction of individuals characterized by relatively low adapt-
ability led to degeneration of a local population. Moreover, apart from the negative effect
of genetic factors, introduction of individuals from a foreign population poses a threat
that some diseases or parasites will be brought to a local population (Dannewitz 2003).
The negative effect of introducing individuals from a foreign population is not lim-
ited exclusively to a population of the same species (Dannewitz 2003, Holmlund and
Hammer 2004).
Conservation and restitution of endangered populations is one of the most essen-
tial aims in fish stocking programmes. At present, the generally accepted procedure is
to obtain stocking material by breeding individuals from a local population and then
to release fish fry to the natural environment, which means that they are mixed with
individuals of a threatened population living in the wild. This procedure is known as
supportive breeding and, in principle, enables us to alleviate the negative consequences
of fish introduction and to increase the total abundance of a population (Wang et al.
2001, Dannewitz 2003).
In fact, the effect of such factors as variable influence of natural selection and mating
or the occurrence of random genetic events during fish breeding lead to differentiation of
reared and wild individuals (Einum and Fleming 2001, Dannewitz 2003, Weir and Grant
2005). Besides, since fish are highly flexible organisms in respect of the phenotype, most
environmental factors that come into play during fish breeding can directly contribute to
the appearance of developmental differences. In a typical hatchery, fish live in conditions
completely different from those in nature, e.g. high fish density in discharge tanks. Thus,
such factors as diet, density, substratum, exposure to predators, or interaction with indi-
viduals of the same species, have a considerable influence on shaping differences between
reared and wild populations (Einum and Fleming 2001, Dannewitz 2003).
Considering the influence of fry stocting on the structure of natural populations, one
must mention the phenomena of inbreeding and genetic drift, which are the factors af-
fecting genetic variation. The intensity of these phenomena is determined by the effec-
tive size of a population Ne, defined as the size of an ideal population that has the same
inbreeding or genetic drift coefficients as the observed population. An ideal population is
characterized by an equal number of males and females, random mating, lack of changes
36
Genetic monitoring of the fish stocking using microsatellite DNA markers
in the abundance of a population between subsequent generations, no overlap of the

generations, random variability in the number of produced offspring, etc. Actually, exist-
ing populations are divergent from an ideal one, thus the effective size of a population
tends to be smaller than the nominal value (Wang et al. 2001, Dannewitz 2003).
Inbreeding is the crossing of individuals more closely related than would appear from
the calculus of probability, should individuals randomly chosen from a whole population
mate (Wang et al. 2001). This type of crossing, especially in natural populations with
a stronger homing instinct (an instinct which drives them home) or spawning in different
seasons, to some extent may have a positive effect. Aggregation of a limited number of
individuals in the same place and at the same time, as a result of the above behavioural
conditions, can considerably increase the chance that related animals will cross, and that
serves as a catalyst shaping and maintaining within a given population a complex of genes
responsible for adaptation to local conditions. In contrast, the incidental nature of genetic
drift, along with selection, migration and mutations, plays an important role in shaping
the genetic variation of a subpopulation of the species (Wang et al. 2001).
The unfavourable effects of inbreeding, however, become fully evident when the size
of a population diminishes. A decline in the abundance of a natural population, caused
by human activities (e.g. due to overexploitation of a population or worse environmental
conditions), which is the main reason why stocking programmes are initiated, implies
a series of phenomena that can threaten the existence of a whole population. A consid-
erably reduced population means that the risk of becoming extinct grows due to some
randomly occurring events. Moreover, a smaller population may experience farfetched
changes on the genetic level. As a result, the population may lose genetic variation,
which enables the population to adapt to changes in the environment, or its adaptability
will be reduced due to the so-called inbreeding depression (Dannewitz 2003). Inbreed-
ing depression is a direct consequence of mating between closely related individuals,
which leads to an increase in the frequency of genotypes versus heterozygotic geno-
types in a population, as a result of which various elements of the adaptability value are
likewise depressed (Wang et al. 2001). This can manifest itself, for example, as loss of
fertility, increased susceptibility to diseases, appearance of developmental abnormalities
or metabolic disorders (Taniguchi 2003).
The influence of inbreeding and genetic drift is not limited to natural populations.
Moreover, in cultured populations, this influence seems to be particularly significant be-
cause it is the main factor responsible for creating differences between populations on
the genetic level (Taniguchi 2003, Weir and Grant 2005). A higher degree of inbreeding
in cultured populations may be caused by many factors, such as: insufficient number of
reproducing animals, an unequal number of mating males and females, selective mating
and other effects leading to a regular selection of specific phenotypes, unequal repre-
sentation of parental genotypes among the offspring introduced to natural populations
or selected for breeding, or else varied survivability of families (Wang et al. 2001).
Supportive breeding, although considered to be the safest form of fish stocking
among all introduction methods, can also be affected by the above phenomena. Just
when some individuals from a population are selected for culture, the risk of a result-
ing depressed genetic variation within a cultured versus natural population increases.
Besides, when the reproductive potential of some part of a population is enhanced by
37
fish breeding techniques, inbreeding and genetic drift appear (while the effective size of
a population decreases) due to an increased variance in the number of offspring in the
whole population (Wang et al. 2001, Dannewitz 2003).
In sum, introduction of individuals from a foreign or cultured population may lead
to farfetched consequences for a natural population occurring on the genetic level, in
general resulting in a lower adaptability coefficient due to introgression. Ecologically
speaking, transfer to a new ecosystem, such as the habitat in which a natural population
dwells, may lead to increased predatory pressure within the ecosystem or appearance
of competition between the introduced and local populations, up to the extent that the
former can replace the latter one (Wang et al. 2001, Dannewitz 2003, Weir and Grant
2005). Furthermore, large changes in the trophic network of the whole biocenosis can
appear, as the introduced individuals exert more pressure on the local plants and other
groups of organisms dwelling in the ecosystem (Holmlund and Hammer 2004).
Applicability of microsatellite DNA as a genetic marker to conservation of ich-

thyological diversity
Our knowledge of complex mechanisms which have an effect on variation of a popu-
lation within a given species is of primary importance for the rational management of
fish resources and conservation of natural fish populations. It is now possible to know
the genetic structure of a population and the influence of the factors which shape it
owing to the up-to-date tools of molecular biology (Sunnucks 2000). Many of the con-
temporary methods of analyzing polymorphism of a nuclear sequence and mitochon-
drial DNA have proven to be applicable to fisheries and aquaculture (Hayes et al. 2005).
Among such methods, analysis of microsatellite DNA is the most popular one, gradually
supplanting other methods (Okumuş i Çiftci 2003).
The structure of microsatellite DNA

Microsatellites, also referred to as short tandem repeats (STRs) are non-coding, in-
dependently segregating fragments of highly polymorphic chromosomal DNA. A single
section of microsatellite DNA is composed of a series of repetitive units, between 1 to 6
base pairs (bp), bordered on both sides by flanking regions (Li et al. 2002). The length
of a whole fragment may range from 20 to a few hundred base pairs (Chistiakov et al.
2006). According to DeWoody and Avise (2000), typical series of repeats are composed
of motifs of a sequence between 2 to 4 nucleotids repeated from 5 to 40 times. The
diagnostic characteristics in studies on the polymorphism of microsatellite DNA are: the
length of a sequence of repeats and a sequence of bases in a given series. Unlike highly
variable repeat series, the flanking regions are evolutionary conservative in closely re-
lated species (Wąs and Wenne 1998). This characteristic makes flanking regions an ideal
site for attaching starters during PCR amplification of microsatellite DNA.
Advantages of microsatellite DNA as a genetic marker

A unique and high degree of polymorphism alongside its widespread occurrence in
a genome, a small size of microsatellite loci and independent segregation make msatDNA
a useful molecular marker. The high degree of polymorphism enables researchers to use
short tandem repeats to analyze the genetic structure on the population level (Wąs and
38
Wenne 1998) or even on the level of a single individual (Artamonova 2007). Additionally,
with the characteristics of msatDNA it is possible to analyze partly damaged genetic mate-
rial, originating from archived samples, such as dry scales or fin rays. Microsatellite DNA
then could be a very useful genetic marker, helping us to reconstruct genotypes or to trace
changes in the genetic structure of a population over some time period.
Evaluation of the genetic variation of a population

The extremely high sensitivity of the method is one of the reasons why analysis of
microsatellite DNA is so popular in investigations on genetic variation and population
diversity. Using this method, a high degree of genetic variation (over 52%) has been
determined among natural populations of trout originating from such geographically
distant parts of the world as France, northern Africa, Greece and the Persian Gulf (Presa
and Guyomard 1997).
Analysis of msatDNA is also an ideal tool for studying the genetic variation on a mi-
crogeographic scale. Numerous studies have enabled researchers to find statistically
significant differences between allopatric and parapatric populations of trout (Salmo
trutta) living in north-western Spain (Sampos et al. 2007), Scandinavia (Östergren 2006),
the southern part of the Balkan Peninsula (Apostolidis et al. 2008), north-western Russia
(Lehtonen et al. 2009) and Poland (Wąs and Wenne 2002).
Statistically significant genetic variation between and within populations has also
been demonstrated for other species of the family Salmonidae, such as Salmo salar
(Säisä i in. 2005), Salmo marmoratus (Fumagalli i in. 2002), Salvelinus alpinus (Bernatch-
ez i in. 2002), Salvelinus fontinalis (Castric i in. 2001), Oncorhynchus kisutch (Olsen i in.
2003) and Oncorhynchus tshawytscha (Banks i Rashbrook 2000).
Using msatDNA has also enabled scientists to evaluate the genetic structure differ-
entiation between two sympatrically occurring forms, i.e. anadromous and sedentary,
of rainbow trout: Oncorhynchus mykiss, (Narum et al. 2004) and trout, Salmo trutta,
(Banks and Rashbrook 2000).
Owing to some specific characteristics of msatDNA like high polymorphism and be-
ing selection neutral, this marker can be treated as an autonomous tool for analysis of
inter- and intra-population variation.
The results of a complex analysis of the microgeographical differentiation of popula-
tions of trout in inland waters in France, completed by Estoup et al. (1998), prove the su-
periority of macrosatellite DNA analysis over the analysis based on allozymes. Much more
often than other genetic markers, microsatellite DNA reveals the occurrence of private
alleles, unique for particular populations or their groups. This is one of the reasons why
msatDNA is a much more successful tool for the evaluation of inter-population variation.
Another benefit of using msatDNA is that it is much better at uncovering linkage disequi-
librium between independent loci. Moreover, unlike allozymes, msatDNA enable us to
estimate genetic variation between age groups of a population. This marker is also more
useful in studies on correlations between genetic variation and geographical distance.
Research has also demonstrated that microsatellite DNA is more helpful in the process of
determination of the source of origin of individuals (the assignment test). The superiority
of microsatellite DNA analysis over analysis of allozymes is also confirmed by studies on
the genetic structure of a population of trout in north-western Spain (Corujo et al. 2004).
39
Natomiast mtDNA jako marker genetyczny cechuje się mniejszą informatywnością

w stosunku do mikrosatelitarnego DNA, o czym świadczą m. in. badania zmian struktury
genetycznej izolowanych populacji troci z obszaru północnej Hiszpanii (Campos i in.
2006). Wyniki zróżnicowania międzypopulacyjnego, otrzymane dzięki zastosowaniu obu
metod, w ogólnych zarysach pokrywały się ze sobą, jednak analiza mikrosatelitarnego
DNA w kilku przypadkach ujawniła istnienie statystycznie istotnych różnic między popu-
lacjami, których nie obserwowano na poziomie mtDNA.
Evaluation of the effect of stocking and consequences of introgression
More evidence for the applicability of msatDNA in the evaluation of the effects of
stocking and consequences of introgression is provided by the results of a study on trout
populations in southern France, carried out in order to assess the genetic consequences of
introducing cultured individuals (Poteaux et al. 1999). The analysis comprised natural pop-
ulations affected by the influence of stocking material (intensive stocking was carried out
annually, or else the introduction had been discontinued a few years earlier), and breeding
strains used for supplementation of both populations. The analysis revealed that although
the same alleles were present in both natural populations, the frequency of alleles shared
with the breeding strains was much higher in the case of a population continued to be
stocked. The results of this analysis made the researchers draw a conclusion that the few
years which had elapsed since the last stockings gave sufficient time for the statistically
significant differentiation to be established between the two populations.
Analysis of msatDNA has also been applied in a Danish study on trout populations
(Hansen et al. 2001). The purpose of this investigation was to assess the degree of the ge-
netic variation reduction due to fish culture, to determine mutual relations between natural
populations and breeding strains and to determine the effect of supportive breeding on the
genetic structure of the examined populations. Natural populations, most probably not ex-
periencing fish stocking, and populations subjected to fish introduction (sometimes rather
intensive) as well as cultured fish strains used for fish stocking were examined. The results
of the analysis showed a varied degree of introgression in fish stocked populations, from
relatively low to moderate. Moreover, the results enabled the researchers to determine in-
trogression in populations until then considered to be genetically intact and enabled them
to distinguish natural populations in which no introgression had occurred, which may then
serve for restitution of endangered local populations (Hansen et al. 2001).
Analysis of polymorphism of enzymatic proteins has proven to be insufficient for
identification of natural and cultured trout populations found in the western part of
the Pyrenees (south of France), and originating from the Atlantic line; the evaluation of
the degree of introgression between these populations was made possible owing to the
analysis of microsatellite DNA (Aurelle and Berrebi 1998).
The advantages of msatDNA have been recognized and used in such studies as, for
example, assessment of the effect of stocking on natural trout populations in many Euro-
pean countries (e.g. Van Houdt et al. 2005, Santos et al. 2006). With msatDNA analyses it
is possible to monitor the current fish stocking programmes (Caudron et al. 2006).
This marker has also been applied to the evaluation of introgressive hybridization
between trout (S. trutta) populations and populations of closely related species like S.
(trutta) macrostigma (Marzano et al. 2003), S. marmoratus (Jug et al. 2005), S. obtu-
sirostris (Razpet et al. 2007) or S. ferox (Duguid et al. 2006).
40
Evaluation of the long-term changes in the genetic structure of populations

Most of the research aiming at the evaluation of stocking on populations focuses on
the current changes in their structure, caused by previous events. This approach ena-
bles scientists to determine the presence of introgression and possibly assess its degree,
but does not allow them to acquire detailed knowledge of the changes in the genetic
structure of a natural population, which are a consequence of this phenomenon, nor
does it enable them to determine precisely the rate of introgression. In order to obtain
this information, it is necessary to know the genetic structure of a natural population in
the time period preceding stocking. However, most often it is impossible to acquire suf-
ficient data on the original population. A possible solution to this problem lies in analyz-
ing archived material, for example dried scales or otholits, collected for dozens of years
at institutes of fisheries, and previously used for determination of the age of individual
fish and a growth pattern.
Analysis of archived material creates many methodological problems. Due to the
character of the material, composed of very small amounts of degraded tissue, allozyme
analysis is impossible. However, dried fish scales and otholits are a valuable source of
DNA necessary for running an analysis of the structure of a population (Nielsen et al.
1999). Unfortunately, the DNA obtained is most often degraded due to the time lapse.
Degradation is most often manifested by the reduction in the amount of DNA and frag-
mentation of DNA threads into small sections measuring a few hundred base pairs or
shorter. One consequence of DNA degradation is low efficiency of amplification of long
fragments of mitochondrial DNA with the PCR method, which basically means that this
method will be unsuccessful. In turn, the small size of loci of microsatellite DNA (about
200-300 base pairs) and its widespread occurrence in a genome make msatDNA an al-
most perfect tool for analysis of archive materials.
With the archived materials, it has been possible to evaluate directly the effect of long-
term stocking on populations of trout in Danish rivers. Analysis of msatDNA obtained
from a collection of dried fish scales has served researchers to trace changes in the genetic
structure of populations over some 80 past years (Hansen et al. 2002). The results of this
and some previous studies, including current samples only (Hansen et al. 2001) have con-
tributed to the elaboration of new strategies of the renewal of populations in which a high
degree of introgressive hybridization has been found. For this purpose, it has been sug-
gested to use supplemental fish stocking using exclusively the individuals identified as be-
longing to a proper local population. The individuals chosen for fish stock culture originate
from the tributaries in which no introduction of a cultured strain has taken place; another
option is to use fish stock material from genetically stable populations despite stocking.
Analogous studies have covered populations of trout in Norway. Likewise, analysis
of msatDNA has been applied to evaluate the effect of long-term introduction on the
stability of the genetic structure of the analyzed populations (Sřnstebř i in. 2008).
The studies on Danish and Norwegian trout populations are model examples of
a broad applicability of analysis of microsatellite DNA obtained from archived material.
It not only allows us to gain detailed knowledge of the changes occurring in a popu-
lation and their rate, but it also makes it possible to identify individuals of the native
population, which is key information for the reconstruction of genetic variation of local
populations (Hansen et al. 2006).
41
Evaluation of modifications in the genetic structure of populations going backwards

dozens of years has also been completed for other species of the family Salmonidae.
Owing to the application of microsatellite DNA it has been possible to assess changes,
occurring since the early 1900s, in the structure of populations of salmon (S. salar) in-
habiting rivers in Scandinavia (Skaala et al. 2006), southern France (Martinez et al. 2001)
and inland waters in Canada (Tessier and Bernatchez 1999).
Fish stocking when planned and carried out inadequately are burdened with a risk
of depressing genetic variation of natural population and destroying their genetic struc-
ture, which may lead to their reduced adaptability and extinction of such populations.
However, modern molecular methods using microsatellite DNA markers enable us to
prepare and control the process of supportive breeding, from selection of individuals
for artificial reproduction up to the evaluation of long-term fish stocking. The examples
presented above allow us to expect that the application of diagnostic methods based on
genetic markers will help to sustain the original level of genetic variation of natural pop-
ulations, to restitute populations vulnerable to extinction and to elaborate programmes
for the conservation of ichthyological diversity.
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44
Hatchery practice in the context
of biodiversity of aquatic
ecosystems
Zdzisław Zakęś1, Krystyna Demska-Zakęś2
1
Department of Aquaculture, Stanisław Sakowicz Inland Fisheries
Institute, 10-719 Olsztyn, ul. Oczapowskiego 10.
e-mail: zakes@infish.com.pl
2
Department of Ichthyology, University of Warmia and Mazury in
Olsztyn, 10-718 Olsztyn, ul. Oczapowskiego 5,
e-mail: krysiadz@uwm.edu.pl
Abstract
The objective of the study has been to determine the effect of hatchery practice on
the biodiversity of aquatic ecosystems. The quantitative and qualitative characterization
of the commercial hatchery practice clearly demonstrates that it is driven not only by
economic profits but also by ecological considerations. The analysis of the fish hatchery
production in the region of Warmia and Mazury (RWM) over the past twenty years sug-
gests that it is becoming more and more diverse, both in terms of the number of taxa
being reproduced and the forms of produced fish stock material. In the 1990s, spawn
of 10 fish species was incubated, whereas more recently the number of reproduced
taxa has increased to 15. Spawn of three species is dominant, i.e. pike (Esox lucius),
vendace (Coregonus albula) and European whitefish (Coregonus lavaretus). While in the
1990s, the spawn of these three species made up 97.9% of the total volume produced,
in the recent years this percentage fell to 86.9%. New species are now hatched such as
rheophilous cyprinid species: vimba (Vimba vimba), barbel (Barbus barbus), asp (Aspius
aspius) as well as some ecologically valuable salmonid fish: lake trout (Salmo trutta m. la-
custris) and brown trout (Salmo trutta m. fario). In some hatchery facilities in the RWM,
work has started on the restitution of Atlantic sturgeon (Acipenser oxyrinchus). Nota-
ble is also substantial technical and technological progress in hatcheries in the RWM
compared to the conditions prevailing in such facilities in the 1990s. Some hatcheries
have been equipped in the most modern equipment and have implemented technolo-
gies which improve the safety and predictability of the production as well as broaden
the species spectrum of fish stock material introduced to open waters in the region of
Warmia and Mazury.
45
Introduction
Man affects the biodiversity of water ecosystems, including the quality and abun-
dance of ichthyofauna, both directly and indirectly. Directly, by organizing commercial
and sport fish catches, poaching and introduction (planned and incidental) of alien fish
species and other aquatic organisms. The indirect influence comprises environmental
pollution, uncontrollable development of water shores, construction of hydrotechnical
facilities or land draining systems. The synergistic effect of the above activities, their long-
term influence and cumulative nature has left its imprint on water ecosystem, reflected
for example by a decline in abundance of many fish species. It is symptomatic that the
first evaluation of the condition of our native ichthyofauna according to the criteria of
the International Union for Conservation of Nature (IUNC 1994) was not completed until
some 10 years ago (Witkowski et al. 1999). The conclusions were not very optimistic,
and the validation which followed showed that the state of our native ichthyofauna had
even deteriorated (Witkowski et al. 2009). It turned out that at least 40% of fish species
were classified as belonging to the most endangered class, i.e. critically endangered
(CR), endangered (EN) and vulnerable (VU) species (Kowalska and Zakęś 2010). Among
these taxa, the dominant are rheophilous fishes, but a highly endangered species cat-
egory was also assigned to some of the so-called lake species, e.g. European whitefish
(Coregonus lavaretus), vendace (Coregonus albula) or European smelt (Osmerus eperla-
nus). In respect of lake fish, the main factor responsible for a decrease in their number is
eutrophication of water bodies, which basically causes worse environmental conditions
in spawning grounds and inferior results of natural reproduction. Passive nature conser-
vation forms (legal sizes and closed seasons) or simple forms of active nature protection
(protecting areas where natural spawning of fish occurs, constructing artificial spawning
grounds) can be successful only in slightly transformed lake ecosystems (Bnińska 1992).
In the case of more eutrophic lakes, it is necessary to apply more effective measures of
active nature protection, for example stocking.
Biodiversity of hatchery production

The forerunners of the active fish species protection involving the stocking of natural
water bodies with material obtained via artificial spawning were obviously ichthyologists
and fishermen. For the sake of clarity, it should be explained that they were not driven
by altruistic motivation only. The main purpose of their work was to maintain or increase
the abundance of populations of the economically valuable fish species up to the level
which would make their commercial exploitation viable. It is worth reminding us here
that the history of fish hatchery practice in Poland is more than a 150 years long (Zakęś
and Jarmołowicz 2009). The species which were first to draw attention were salmonids,
including European whitefish. In the second half of the 20th century an important role was
assigned to pike (Esox lucius) and vendace. The sex products were obtained mainly from
wild fish captured from natural waters either during the pre-spawning or the spawning
period. It should be emphasized that in a situation when, due to progressing eutrophica-
tion, the conditions prevailing in spawning grounds deteriorate but the water bodies are
still suitable for the growth of a given species, it is recommendable to carry out catches of
wild spawners, obtain sex products, incubate spawn and stock open water bodies with the
46
Hatchery practice in the context of biodiversity of aquatic ecosystems
produced material. Such activities are both economically and ecologically viable, especially
in respect of species threatened with extinction. In Poland, as a rule wild spawners of four
species of fish, i.e. pike, vendace, European whitefish and pikeperch (Sander lucioperca),
are obtained. In the 1990s, the spawn of the first former species made up 93.6% of the
global amount of spawn
incubated in Polish hatch-
eries. The dominant was
vendace spawn, which
constituted 49.9% of the
total quantity (Zakęś and
Falkowski 1999). Most of
the fish stock material was
produced in Warmia and
Mazury. Of the total annual
average volume of spawn,
i.e. 6,737 l, about 4,466 l (ca
66% of the global amount)
was incubated in Warmia
and Mazury. With respect
to species, 78% of the to-
tal countrywide amount
of pike spawn, 66.7% of
vendace and 49.5% of Eu-
ropean whitefish were incu-
bated in this region (Zakęś
and Falkowski 1999). In the
hatcheries located in Warm-
ia and Mazury, the spawn of
vendace, pike and European
whitefish made up 97.9%
of the total amount (Fig.
1a) (Zakęś and Jarmołowicz
2009). Thus, the region of Figure 1. The annual average percentage of spawn produced in the
Warmia and Mazury domi- region of Warmia and Mazury, (a) the 1990s, (b) recent years
nates in Poland, both in terms of the amount of incubated spawn and the number of
hatchery facilities; it is also characterized by the highest lake density (Zakęś and Falkowski
1999, Zdanowski et al. 2009). For this reason, in the further part of this paper we shall rely
on the data concerning this part of Poland. Back in the 1990s, although the three afore-
mentioned species were dominant, the region of Warmia and Mazury was distinguished by
the most species diverse hatchery production. At that time, the hatcheries in Warmia and
Mazurt incubated spawn of 10 species, whereas in Wielkopolska only eight species were
hatched and in Pomerania no more than three (Zakęś and Jarmołowicz 2009). Our analysis
of the data from the last years has demonstrated that the species diversity in the hatchery
production in Mazury increased. Recently, spawn of 15 species has been incubated in the
47
hatcheries situated in this region (Fig. 1b) (Zakęś and Jarmołowicz 2009). Although the
spawn of three species, that is pike, vendace and European whitefish, continues to prevail
(it is now equal 86.9% of the total volume of incubated spawn), noteworthy is the fact that
the volume of spawn of other species has considerably increased, from 2.1% in the 1990s
to 13.1% in the recent years (Zakęś and Falkowski 1999, Zakęś and Jarmołowicz 2009). It is
not only the species diversity of the fish hatching production (the number of hatched taxa)
that has improved but likewise the volume of production of “new” species. Moreover, it
should be noted that in the last two decades, the absolute amount of incubated spawn in
the region of Warmia and Mazury has not changed considerably and is now estimated to
equal 4350 l annually (Zakęś and Jarmołowicz 2009). But the hatcheries now also produce
spawn of rheophilous cyprinids: vimba (Vimba vimba), barbel (Barbus barbus), asp (Aspius
aspius) as well as some ecologically valuable salmonids, such as lake trout (Salmo trutta
m. lacustris) and brown trout (Salmo trutta m. fario) (Fig. 1b). The family Acipenseridae is
also worth mentioning, especially Atlantic sturgeon (Acipenser oxyrinchus). All these facts
indicate that the species diversity of the fish hatchery production is advancing.
Most of the spawn incubated in the hatcheries in Warmia and Mazury is still ob-
tained from wild spawners, particularly pike, vendace and European whitefish. In the
recent years, catches of wild fish for spawning in the water bodies in Warmia and Ma-
zury have shown a rising tendency, e.g. in 2006 about 41 tons were obtained, whereas
in 2008 there were 66 tons of fish captured (Zakęś and Danilewicz 2010). Basically,
catches of pike spawners have increased, as their share by weight is now 75% of the to-
tal catches of spawners. In turn, the amount of captured vendace has declined from 12
tons (2006) to 10 tons (2008). Analogously, the catches of European whitefish spawners
have diminished. It should be added that the fish catches limits for these species set up
by the Marshall Office of the Province of Warmia and Mazury show an increasing ten-
dency in the recent years, whereas the amount of performed catches is characterized by
a decreasing trend; in respect of vendace, they covered just 40% of the assigned limit
and 27% regarding European whitefish (Zakęś and Danilewicz 2010). Unfortunately, this
decrease in the catches of wild spawners of coregonid fish can be attributed mainly to
a decrease in the abundance of natural populations of these fish species. As a result, the
fish hatchery production in the recent years has been dominated by pike, which now
makes up as much as 51% of the total spawn incubated in Warmia and Mazury (Fig. 1b)
(Zakęś and Jarmołowicz 2009). Regarding the species whose abundance is drastically
decreasing and the catches of wild spawners are small, the only effective method of
preserving their gene pool and producing fish stock material at a level which will enable
us to maintain or to improve their ecological status is by creating and keeping spawners
under controlled conditions. Such activities have been undertaken at science institutes
and fish farms to preserve European whitefish (Szczepkowski 2009). Similar work has
been carried out for a few years on Atlantic sturgeon and noteworthy if the fact that
the driving aim is the restitution of this species (Kolman 2009). Also, a method has been
elaborated for keeping pikeperch spawners in recirculation aquaculture systems (RAS)
(Zakęś 2009). The above list of the activities pursued in hatcheries is probably the best
confirmation of their positive and unquestionable role in maintaining biodiversity of
natural water bodies. The major purpose of these activities is producing fish stock mate-
48
rial of the aforementioned species, becoming independent from catches of wild spawn-
ers and stabilizing fish stocking treatments of open water bodies.
The technical infrastructure of hatcheries versus hatchery production structure

Clearly seen progress in the Polish fish hatchery has taken place over the last several
years. The progress needs to be considered in terms of technical infrastructure. Ab-
sorption of means from the European structural fund, under the Sectoral Operational
Programme Fisheries and Fish Processing 2004-2006 has contributed to the improve-
ment of the technical condition of some hatcheries. It is assumed that at least a similar
effect will be brought about by the following Operational Programme Sustainable
Development of the Fisheries Sector and Fisheries Areas 2007-2013. Although it is
not possible to speak of general improvement of the hatchery industry facilities, it
should be stressed that any comparison of the infrastructure back in the 1990s and
the recent years shows a considerable progress (Zakęś and Falkowski 1999, Zakęś and
Jarmołowicz 2009). Some of the hatcheries now have the most modern equipment
and use the technologies which increase the safety and predictability of production,
thus improving its effectiveness. In the 1990s none of the hatcheries in Warmia and
Mazury had technologically advanced equipment for water treatment, e.g. oxygen
generators or microsieves. At present, 8 hatcheries are equipped with oxygen genera-
tors and 9 have microsieves (Zakęś and Jarmołowicz 2009). Doubtless, this is a certain
indicator of the development in the fish hatchery industry in the region. However, it
should be clarified that quite many hatcheries (ca 25%) still lack basic tool which en-
sure the effectiveness of hatchery production, i.e. the thermoregulation system. The
hatchery potential production capacity over the last twenty years, as measured by
the number of incubation apparatuses in the region of Warmia and Mazury, has not
changed (there are 2,300 incubators; Zakęś and Jarmołowicz 2009). Nonetheless, for
some time now we have been observing a very beneficial development in that new or
refurbished hatcheries diversify their production by adding new, the so-called ‘more
difficult’ species, which are often characterized by high ecological value. They are the
taxa whose successful spawning is conditioned by subjecting spawners to thermal
stimulation; sometimes it is also necessary to apply hormonal injections stimulating
maturation, ovulation and spermation. It is also worth noticing that our knowledge
about fishes, that is controlled fish reproduction and environmental requirements,
has largely improved. Let us remind ourselves that until recently fish hatching, in both
cognitive and practical aspects, focused mainly on species which were of great com-
mercial importance. Our knowledge on hormonal and environmental stimulation of
reproduction, short- and long-term storage of semen, unsticking and incubation of
spawn of the economically important fish species has proven to be useful for under-
taking attempts to reproduce new species, which until now have been of no interest
to commercial hatcheries (Zakęś et al. 2010). What matters is that in many cases the
results of scientific research have found practical application. All the activities which
aim mainly at elaborating an effective method for production of fish stock material of
a particular species are an inherent part of the work the purpose of which is to pre-
serve ichthyofauna and maintain biodiversity of our waters.
49
It should also be no-

ticed that the range of the
production profile in many
hatcheries has been con-
siderably extended. Until
recently, they were typical
fish hatcheries in which
spawn obtained from wild
spawners was incubated,
mainly under field condi-
tions. Hatched larvae were
mostly used to stock open
water bodies. Only a small
portion of larvae was reared Figure 2. The annual average share of species reared in hatchery
in carp ponds or in the so- of Warmia and Mazury over the last years.
called off-lake ponds (Zakęś and Falkowski 1999). At present, there are a few hatcher-
ies possessing rearing facilities with recirculation aquaculture systems. In the 1990s,
rearing hatched larvae in a RAS was carried out in just 2 hatcheries, but in more recent
years, nearly every other hatchery has such systems (Zakęś and Jarmołowicz 2009). In
fact, calling such facilities ‘a hatchery’ is not quite correct. We should rather name them
hatching and rearing facili-
ties (H&RFs) (Figs. 3 and 4).
The total annual amount
of reared material at the
H&RFs may not be very im-
pressive (ca 2.7 million in-
dividuals), but noteworthy
is the rich species compo-
sition of the reared mate-
rial, which encompasses
Figure 3. A new hatchery and rearing facility at the Inland Fisheries
14 taxa (Fig. 2). Among the
Institute in Olsztyn (photo: M. Szczepkowski). reared species, ecologically
valuable fish dominate, i.e.
European whitefish (550
thousand indiv.), pike (415
thousand indiv.), brown
trout (250 thousand indiv.)
or European eel (200 thou-
sand indiv.). The acquisi-
tion and implementation
of methods for rearing the
fish stock material at the
H&RFs makes it possible
Figure 4. The rearing room (inside) in the new hatchery and rearing facil-
ity of the Inland Fisheries Institute in Olsztyn (photo: Z. Zakęś). to stock open waters dur-
50
ing the seasons which are environmentally most suitable for fish. This can be of much
importance for coregonid fish, which are stocked in an unfavourable season, e.g. when
lakes are under ice cover. It should also be emphasized that larvae are reared not only
to the fingerlings, but some of the material is kept in RAS until they become spawners.
Having cultured spawners makes the production of fish stock material more predictable
but poses several challenges in front of ichthyologists. One such challenge is, for exam-
ple, the elaboration of a feeding method that will ensure production of good quality
sex products. It should also be remembered that appropriate genetic diversity need to
be maintained. Creating basic biotechniques of rearing and keeping of fish spawners
is important both economically and ecologically. It is actually a form of ex situ species
conservation.
Conclusions
The primary aim of commercial fish hatcheries was production of fish stock material
(mainly larvae) and maintaining the abundance of economically valuable species of fish
in natural ecosystems. Until recently, this activity had mainly an economic aspect whereas
its ecological importance was more of an incidental character. Fishing companies operat-
ing on open waters were first of all interested in species most vulnerable to unfavorable
changes in the aquatic environment, caused by eutrophication of water bodies, for ex-
ample the family of coregonid fish. Simple forms of active conservation of these species,
such as catches of spawners, artificial reproduction, incubation of spawn in hatcheries
an stocking open waters with hatched fry have contributed to the maintenance and pos-
sibly improvement of the status of these species. Actually, we are unable to say what
status they would have now without regular stocking treatments done in the past years
using fish stock material from hatcheries. However, we feel that the knowledge gained
over dozens of years of running hatchery business cannot be overvalued. The information
bank on biology, reproduction, physiology and behavior of fish, initially of high value to
commercial fish, now enables us to successfully reproduce and produce stock material of
many fish species, aggregating tens of taxa. The hatchery industry profits greatly from
the development of techniques and technology, to which people involved in aquaculture
have also made a large contribution. In sum, hatchery production is becoming more and
more diverse, in terms of the number of reproduced taxa and forms of produced fish stock
material (from larvae to spawners). However, considering the fact that a real progress in
the hatchery industry has occurred in the last ten years, many reproduction techniques or
technologies applied to fish stock material production have not been completely perfected
yet. Some are in the initial, experimental phase, especially the ones which deal with new,
more difficult species. Working on reproduction and fish stock material production meth-
ods or keeping cultured spawners is certainly an important tool to be used in programmes
of conservation of particular taxa, whose main objective is the preservation of biodiversity
of our ecosystems. Using only passive nature conservation forms, particularly in strongly
eutrophic lake ecosystems, although necessary, is not fully successful. In turn, active na-
ture conservation programmes can be effective only is complementary in character and
combine activities pursued by lake fishermen, science institutes and organs of the state
administration. It is also important to work out legal regulations on fisheries management
51
and species conservation of ichthyofauna, in the context pf activities favoring the mainte-
nance of biodiversity of aquatic ecosystems.
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52
Fish stocking versus biodiversity:
peaceful co-existence or
continual conflict?
Jacek Sadowski, Jolanta Kempter
West Pomeranian University of Technology in Szczecin,

e-mail: jacek.sadowski@zut.edu.pl
Abstract
Fish stocking, being one of the fishery management treatments carried out by users
of lakes, rivers and other water reservoirs, means that a new gene pool is introduced to
a given water body. Such changes can vary from very small ones, such as supplementa-
tion of an existing population of a species with fish originating from outside the water
body, up to an introduction of a completely new species, which has not been present
there before, including an introduction of a species outside its natural range of occur-
rence. In each case, fish releases can affect the broadly understood biodiversity of organ-
isms inhabiting a given water body. This paper contains a presentation of the historical
approach to the question of fish brood stocking and the influence of the binding legal
regulations on decisions made by users of water bodies as part of their own fish release
policy. It also discusses such a selection of spawners for reproduction that is rational
from the point of view of the conservation of genetic variation. Examples from Poland
and other countries have been presented to show what negative effects on aquatic
ecosystems can appear due to bad fish stocking management. The recapitulation of the
paper contains an evaluation of the current situation in the fish stocking management
and its possible influence on the conservation of biodiversity in Polish waters. Questions
are also raised on how fish releases should be conducted in order to sustain the biodi-
versity of aquatic organisms.
Introduction
The concept of biodiversity – biological diversity – was for the first time used by Tho-
mas Lovejoy in 1980. Since then, it has made a spectacular career, especially in the context
of extinction of species and threats to the natural environment. Since the Earth Summit
held in Rio de Janeiro in 1992 and the enactment of the Convention on Biological Diversity,
biodiversity has been understood as the diversity of all living organisms on the Earth, in
terrestrial, marine and freshwater ecosystems as well as in ecological assemblages which
53
they are part of. Biodiversity concerns the diversity within a species (genetic diversity), be-
tween species and the diversity of ecosystems. Biodiversity is often used as a term defining
the sum of species or ecosystems in the analyzed or compared areas. There are now many
definitions of biodiversity. It can even be claimed that unlike the actual biodiversity, which
is continually diminishing, the variety of its definitions keeps growing. Many indices are
applied to assess biological diversity, of which the most often applied ones are: species
richness, rare species richness, Margaleff, Pielou, Shannon and Simpson indices.
The fish stocking management in the context of biodiversity

The history of fish stock releases is longer than that of the concept of biodiversity. It
can certainly be claimed that the history of releasing fish brood is as long as the history
of man’s civilization and fisheries. A perfect example is “our” carp, whose “successes” in
the environments foreign to this fish species have become proverbial and today carp in-
habits practically all conti-
nents (Balon 1995). Its last,
although doubtful, success
was the colonization of
Australia (Smith 2005) and
New Zealand (McDowall
1979) in the 20th century
(Fig. 1).
Fish stocking is intrinsi-
cally linked to the questions
of acclimation and intro-
duction, which over many
years have made a great
career in the fisheries sci-
ence. Both phenomena, if
they take place in what is
commonly known as open
waters (natural water bod-
ies), can clearly add to the
diminishing of biodiversity.
An excellent source of infor-
mation on such effects, at Fig. 1. An illustration presenting the situation of carp in Australia
– any semblance to a rabbit is intentional (Roberts and Tilzey 1997).
least recorded until 1952,
is a book written by Sakowicz (1952), who demonstrated that the two superpowers of
that time had done enormous “work” in this field. The genetic mess that the Soviet or
American fisheries had made was never repeated on such a big scale in later years. Could
it be so because Watson and Crick discovered DNA? However, even after their discovery,
some spectacular reintroductions of fish have been carried out, without paying any at-
tention of gene pools. At present, the top position in such undertakings is held by China,
which has already acclimatized species from different parts of the world, e.g. the Nile
tilapia (Oreochromis niloticus), rainbow trout (Oncorhynchys mykiss), largemouth bass
54
Fish stocking versus biodiversity: peaceful co-existence or continual conflict?
(Micropterus salmonides) or turbot (Psetta maxima) (Li 2003). Although these fish have
been imported to keep in aquaculture facilities, it is highly probable that they will colo-
nize the Chinese waters permanently because of the widespread aquaculture practice
and the methods it applies in China, such as using natural water bodies (cage and pen
culture). Recently, possible acclimation of iridescent shark (Pangasianodon hypophthal-
mus) in North America has been discussed (McGee 2010). Another example is the ac-
climation of the Nile perch (Lates niloticus) in Lake Victoria and its adverse effect on the
abundance of local populations of cichlids (Ogutu-Ohwayo 1990, Witte 1992a, 1992b),
or carp in Australia (Smith 2005) as well as prussian carp (Carassius auratus gibelio) in
Poland, where it competes strongly with crucian carp (Carassius carassius).
On the other hand, introduction of new species in waters in which they have not dwelt
before may contribute to an increase in the total number of species inhabiting a given
area. Poland, where widespread acclimation of fish has brought about an increase in the
total number of fish species without changing the number of indigenous ones, is an ex-
ample that is closest to us. Although some of those fish stock releases can be regarded
as harmful ones to the fishery economy, such as the fish releases carried out by Max von
Borne, they have led to the acclimation, by some considered undesirable, of brown bull-
head (Ictalurus nebulosus) (Copp et al. 2005), pumpkinseed sunfish (Lepomis gibbosus)
(Przybylski 2006) or the crayfish Orconectes limosus (Grabowski et al. 2005). These species
are regarded as fish pests, but if not for the presence of Orconectes limosus in our waters,
many people in Poland would not have seen a live representative of crayfish. This does not
mean, of course, that a way of raising biodiversity is unlimited stocking of water bodies
with alien species, but the result of such releases does not have to be always negative.
In Poland, fish releases are carried out for a number of purposes, e.g. restitution
and conservation of rare fish species, absurd rules regulating leases of fishing territo-
ries and, to the least extent, economic considerations. There are also planned angling
fish stock releases and, unfortunately, some social actions consisting of fish releases to
“ownerless” water bodies. In most cases, such activities should be deemed harmful to
the diversity of ichthyofauna. There are several reasons. First of all, there is not enough
fish stock material of good quality in comparison to the fish brood releases declared
by fishing grounds users, which makes them fight for fish stock material regardless its
quality and origin. In this respect, the most spectacular is the fish stocking of lakes with
European whitefish. Although no one will admit publicly that this is a right treatment,
it is broadly applied. On the one hand, there is the pressure of fishing grounds manag-
ers to fulfill fishery plans. On the other hand, no one actually controls what goes on in
fishery regions. As a result, there are many fish brood releases that are uncontrolled, ir-
rational and absurd, in terms of both biology and economy. The underlying cause is the
binding regulations which specify the rules of competitions for leasing fishery grounds
and a resulting ‘”rat race” between competitors. Fortunately, an amendment to the Act
on Inland Fishery (Journal of Law, 2010, no 200, item 1322), which makes a provision
for extending a lease contract without having to run another competition, gives us some
hope that this “drive” to carry out fish stock releases will at least partly be restrained.
Unfortunately, there are also scientific reports on irrational activities, whose results
are difficult to envisage and motivation is hard to comprehend. An example could be
55
the introduction of salmonid hybrids to open waters (Pilecka-Rapacz et al. 2010) or free
creation of acipenserid hybrids (Kolman and Szczepkowski 2001). In the latter case, the
hybrids are at least not introduced to natural waters. When the Act on Inland Fishery
was amended, certain legal restrictions were introduced on the then uncontrollable in-
troduction of alien species and hybrids to the natural environment, but the up-to-date
practice seems to suggest that this law will not be successfully enforced, showing that
paper is more patient than man.
In respect of marine fish, there are negative examples of managing fish stock re-
leases, too. We were very close at one point to fish stocking the Polish zone of the Baltic
Sea with cod. Considering the earlier Danish experience (Střttrup et al. 2005; 2008a;
2008b), the costs of such an undertaking would be huge and there would be a real risk
of depleting the gene pool. In fact, that would have been an attempt at reaching some
current political aim with the taxpayers’ money. It is much more important to manage
the existing resources rationally and to successfully protect the habitats and spawning
grounds of cod. This should suffice for the conservation of this species.
Objectively speaking, does it make sense to carry out fish releases in Poland? With
respect to economy, the answer is unfortunately ‘no’. However, for saving some spe-
cies and sustaining diversity within a given species – provided the genetic resources are
adequately managed – the answer is certainly positive. But than more outlays should be
allocated to monitoring fish stocking, including the control of genetic resources. There-
fore, a group of fish should be selected for which such management will be run.
Nonetheless, it is possible to find and list positive examples. On the global scale, we can
point to the management of migratory fish populations in the Pacific Ocean. Despite some
errors made, it can be firmly stated that without the widespread fish stock releases carried
out by the United States, Canada and Japan, populations of Pacific salmon would now be
totally regressing. This does not mean that their situation is now completely under control.
Another positive example is the campaign for the sake of restituting Atlantic salmon in
Polish waters. Although the Atlantic salmon fish living in Polish waters are offspring of the
stock material imported in relatively small quantities from the Daugava River in Latvia, we
can claim that this has been a successful reintroduction (Bernaś et al. 2009).
Another example is the management of deep-water fish in Poland. With respect
to vendace, running the fishery economy based on artificial spawning and fish brood
release enables fishermen to achieve profits, but these activities most probably cause
a decline in the gene pool of this species. As regards European whitefish (Coregonus
lavaretus), its natural reproduction has been severely restricted by degradation of the
natural environment, therefore conservation this species requires artificial reproduction.
Because European whitfish is one of the species most frequently introduced to open
waters in Central Europe, its gene pool is heavily mixed, although some recent research
has proven that there is quite large genetic variability between particular populations
(Kohlmann et al. 2005, Kempter et al. 2010). In 1782, when Bloch first described the
whitefish in Miedwie Lake, he could not expect that this would become one of the most
fertile species in creating diverse whitefish populations and that stocking lakes with
this fish would enhance the diversity of whitefish. In the territory of today’s Poland,
whitefish stock releases certainly began back in the 19th century. The following white-
56
fish were used for releases: whitefish from Miedwie Lake, whitefish from Holstein and
densly-raked whitefish from Pejpus Lake. Releases of fish stock material obtained from
these populations were definitely conducted before World War One (Sakowicz 1952, Kaj
1955). Afterwards, they grew in scale and in practice were not submitted to any genetic
monitoring. This has not changed until today, despite some attempts to control fish
stocking. Based on research data, we can presume that today the following whitefish
populations exist: populations of whitefish from Miedwie Lake (Sadowski et al. 2010),
migratory whitefish from the Bay of Puck (Heese 1985), migratory whitefish from Łebsko
Lake (Heese et al. 1999; Fopp-Bayat and Wiśniewska 2010) and noble densely-raked
whitefish (Falkowski and Wołos 2007, Szczepkowski et al. 2010), close to the whitefish
described by Peters in Pełcz Wielki Lake (Peters 1874). It is important to make a decision
which populations and which lakes should be distinguished and considered as “mater-
nal” ones for the whole population. In this case, it will be necessary to coordinate the
work of fishers, researchers and state officials in order to maintain the biodiversity of
whitefish within the species.
With respect to vendace, due to the fish stocking practice described above, com-
pletely free fish releases with this species should be allowed provided its high interspe-
cies homology. The question of fish releases of other species, which have been inten-
sively stocked, such as pike-perch, pike, tench or crucian carp, yields a similar answer
(Kempter et al. 2011).
Another problem, which must not be overseen, and which in practice is beyond
any control, is the question of transmission of pathogens, accompanying fish brood
releases. Viruses should be mentioned first. The “career” of both VHS and KHV is suf-
ficient evidence. The known viruses can be diagnosed before introducing fish to a new
environment, but in practice there is no method to protect us from introducing a virus
unknown to science.
By analogy, transfer of fish stock material can cause transfer of parasites, although
these seem more easily controllable. The transfer of parasites occurs both within and
outside their natural range of occurrence. An example could be the nematode Anguil-
licola crassus, widespread in European populations of eel (Anguilla anguilla) but origi-
nating from Asia (Własow 1991, Rolbiecki et al. 2000). It is obviously a parasite that
threatens the sustenance of eel in European waters.
Local populations and their genetic differentiation

Genetic characterization of the diversity within local populations has been studied
in many species. In particular, molecular differences between populations have been
examined in salmon (Verspor 1997, Ayllon et al. 2006, Lage and Kornfield 2006), cod
(Nelson et al. 1994, Arnason and Palsson 1996) and sturgeon (Ludwig et al. 2004, Kul-
lander 2004). The need to introduce protection programmes for these fish meant that
it was necessary to create a molecular data base which would enable evaluation of the
degree of their differentiation. Considering the fact that research teams working on rare
or threatened species focus on searching of the best possible ways of preserving genetic
variation within populations, one should examine the applicability of commonly used
molecular markers and assess to what extent they can fail. The gene pool of a population
57
which enables its identification is a consequence of the morphological or geographical

differences of the individuals which create that population. Because there are many at-
tempts to verify diversity within populations, in ichthyology, genes located in the mito-
chondrial or nuclear DNA are examined. At present, particularly noteworthy is the gene
responsible for cytochrome b (cyt.b), NADH dehydrogenase coding gene (ND-1) or the
cytochrome oxidase gene (Cox-1). These genes have been used successfully to differenti-
ate populations of whitefish in order to find the haplotype closest to the whitefish from
Miedwie Lake (Kohlmann et al. 2005, Kempter et al. 2010). A sequence of a fragment of
cyt.b gene has been applied to analyze inter-population diversity within whitefish (Os-
tbye et al. 2005), Coilia ectens (Cheng and Lu 2005), Coilia mystus (Cheng et al. 2008),
beluga sturgeon (Huso huso) (Dudu et al. 2009), and sockeye (Oncorhynchus nerka)
(Bickham et al. 1995). Designing starters which facilitate amplification of a fragment of
Cox-1 gene has contributed to using the conserved region for identification of popula-
tions among sterlet sturgeons (Hofsoe 2010, Neumann 2010) or eels (Trautner 2006).
Abundance of a population versus its being threatened

Reproduction of domesticated animals is said to be critical if the total number of
females is less than 100 or the total number of males is less or equal 5 or the total popu-
lation encompasses fewer than 120 individuals and the share of animals ready to repro-
duce in this population is about 80%. A threatened population is defined as a group of
individuals, in which the number of mature females is more than 100 but fewer than
1,000 or the total number of mature males is fewer or equal 20 but more than 5, or the
total number of individuals in a population is more than 80 but fewer than 100 (Słomski
2010). The above categories identify critically threatened or threatened populations, for
which in situ protection programmes or programmes based on cultured stock material
are worked out. Taking into account the amounts of material used for fish reproduction
and for creation of spawning herds of threatened species, the genetic step and inbreed-
ing in the cultured population of fish should be considered. Such studies, however, are
rarely performed and the material (including hybrids) is released to open waters. An
example could be the hybrids of sturgeons (Kolman and Szczepkowski 2001), which al-
legedly do not spawn naturally with other species of the family in open waters. However,
when such a hybrid spawner is caught and its spawn obtained, a result could be the
transfer of traits of the hybrid and pollution of the gene pool of a pure species.
A critically low number of spawners has been frequently taken advantage of in repro-
duction of sturgeons (Kolman and Lutikov 1998, Kolman et al. 1999). Selection of spawn-
ers which will ensure maintaining adequate genetic variation should rely on keeping the
measure of adjustment of a population (which is the stock material) in the sense of surviv-
ability of individuals until they start spawning and on restraining the inbreeding depres-
sion. Formation of spawning herds, which will be a source of a gene pool for subsequent
generations, ought to be based on calculations of their minimum number (the so-called
number of effective population Ne). Determination of the minimum number of individu-
als will guarantee high survivability of the produced flock, and will also ensure that once
the brood material is released into open waters, it will be able to successfully compete
for survival. Obviously, when spawners are caught, it is impossible to obtain 100 or 150
58
individuals, and reproduction based on 1 or 2 females and a few males is dangerous. As

Fopp-Bayat et al. (2010) state, there is a risk that the number of individuals in a popula-
tion will be reduced to a dangerous threshold such as 30 specimens. This moment should
alert the breeder, who is then obliged to start selecting a new breeding herd. As for genes
which are widespread, and included in fishing plans, new breeding herds can be created.
But it is much more difficult when this programme comprises a threatened species and
sites where new fish for reproduction can be obtained are very few.
The species imported to Poland, including the American paddlefish, are also subjected
to artificial reproduction treatments based on an insufficient number of spawners (Kolman
et al. 2004). Particularly worrying is the case of the American sturgeon, which is now being
reintroduced to Polish waters. Fertilized eggs obtained from 2 females and milt produced
by 7 males were brought to Poland in 2005. Further material (fertilized spawn), imported
in 2008, originated from artificial reproduction involving 3 females and 5 males (Kolman
and Wiszniewski 2008). A question arises whether a population produced from such few
individuals, and intended to populate Polish rivers, will not be burdened with an exces-
sively high degree of inbreeding? If it will not experience such phenomena connected with
inbreeding depression as depressed fecundity of reproducing fish, considerably increased
mortality of early developmental stages or a higher incidence of revealed developmental
anomalies? But, on the other hand, is there another solution?
The situation is different in the case of European whitfish. The search for a pure
haplotype of the historical whitefish from Miedwie Lake has proved to be more difficult
than excepted. The fish for the research were obtained from the lakes Miedwie, Ińsko,
Wisola, Marianów, Woświn, Morzycko, Czarne and Śremskie (in total 69 individuals). The
analyses proved that the sequence most closely resembling the reference ones appears
in the individuals captured in Śremskie Lake. The whitefish caught in Miedwie Lake rep-
resented two haplotypes (H3 and H5), also found in Morzycko Lake (H3), Ińsko, Wisola
and Śremskie lakes (H5) (Kohlmann et al. 2005). The problem we face here, i.e. lack of
distinct genetic differentiation of whitefish from Miedwie Lake, is caused inter alies by
fish stock releases. All the above lakes, except Czarne Lake (which lies in Drawieński
National Park), have been stocked with fish. The presence of haplotypes, which con-
firm that the lakes are populated by open hybrids of whitefish and vendace or peled,
indicates that such individuals have escaped from hatcheries to open waters. It was not
until recently that studies demonstrated that Miedwie Lake whitefish had survived in
that lake (Sadowski et al. 2010) and the sequence presumed to be a reference one and
published in the NCBI (AB034824) corresponds to the haplotypes changed due to ac-
climation (Miedwie-the Czech Republic-Japan).
Molecular markers used in studies on rare and protected species

Until the present day, many molecular markers have been applied to determine the
stability of analyzed taxonomic structures (herds, populations or species). Such evalua-
tion is possible via analysis of loci, RFLP technique as well as analysis of quickly evolving
molecular systems, including mitochondrial DNA. A measure testing the condition of
a taxon is the evaluation of the variation between the individuals of which it is com-
posed. Although the abundance of populations of rare or threatened animals is usually
59
very low, it has been found out that in many cases they do not represent a low level of
genetic variation. This means that we can expect a situation in which a species that has
maintained “normal” genetic variation will become extinct. How can we, therefore, as-
sess the risk and determine the threshold danger? We can be aided by good knowledge
of the evolution rate of a given taxon as well as an average rate of mutations occurring
on the level of mitochondrial DNA. These two features are strongly correlated with the
history of a given species and, to be more precise, with the nature of a decrease in its
abundance over the past tens of years. Reduction of spawning herds certainly has an
effect on possible gene exchange and consequently on the number of possible combina-
tions during crossing. In 1989, Tajima’s neutrality test was worked out, which enables
us to compare the number of segregating sites in an analyzed sequence per one segre-
gation site. A negative value produced by the test shows that the analyzed population
is genetically regressing and needs active protection. It is difficult to asses, anyway, to
what extent and since when the loss of genetic variation is harmful to a population or
a species. Thus, it is proposed to assess the degree of genetic variation based on more
than one class of molecular markers because otherwise the data may not be representa-
tive to the whole genome (Avise 2008). We should therefore consider whether the pro-
tection plans actually elaborated for the species listed on the Red List of Threatened and
Endangered Animals in Poland (Głowaciński 2002) should not rely, in equal measures,
on environmental and behavioural factors, which – like molecular traits – can disrupt the
survival of a population or a species, or even make it impossible.
Effect of fishery management on genes, species and ecosystems

A) Positive influence:
1. maintaining fish resources by stocking flowing waters, which are at risk of becom-
ing extinct due to hydrotechnical constructions (dammed reservoirs, dams) blocking
the flow of rivers,
2. contributing to the sustenance or reintroduction of threatened species by produc-
ing fish brood material obtained from spawners living in the wild (huchen, lake trout,
burbot or wild carp).
B) Negative influence;:
1. decreased heterozygosity of fish resources due to fish stock releases,
2. quantitative and qualitative changes in ichthyofauna caused by fish stocking,
3. changes in the chemical structure of water (ichthyo-eutrophication),
4. transfer of fish diseases due to fish stocking of the material that has not under-
gone complex tests (against viruses, bacteria and parasites),
5. overfishing.
Summary
The conflict between those who want to preserve biodiversity and the fishery indus-
try will continue, for a number of reasons. On the one hand, we will witness irrespon-
sible fish releases, stimulated by some particular interest; on the other hand, we can-
not exclude some expensive and spectacular actions undertaken to preserve “imagined
60
biodiversity” (especially when public funds are “easily” at hand). The latter development
can be illustrated by quoting an example of fire fighting water tanks in Szczecin being
assigned a status of Natura 2000 habitat area because of newts which dwell there (JoL
2010 No 77 item 510). This is pure nonsense but ... In conclusion, the number of species
and populations which will be saved when fish releases are stopped or, on the contrary,
continued rationally, will depend exclusively on the outcome of the activities carried out
by persons involved in the fisheries and in environmental protection. No administrative
regulations and procedures will be able to prevent changes triggered by irrational hu-
man actions. To find the golden mean, we should clearly define the guidelines for both
fish releases and nature conservation for the sake of preserving biodiversity. Consensus
seems most easily attainable when dealing with critically endangered species. Lengthy
discussions whether a species or its local population should be protected might end
when both disappear (like vimba in Polish rivers). Not wishing to suggest answers, we
leave the reader to make judgment.
Back in 1952, Professor Sakowicz wrote about the need to change the composition
of commercial ichthyofauna by introducing more valuable species. Today, this form of
fishery management seems unreasonable. It is more because of the high costs of such
an approach that valuable species are possibly helped to survive in a given water body,
sometimes via fish releases.
From the point of view of the inter-population biodiversity conservation, it seems right
to limit fish releases as a fishery management treatment. A decision to run fish stock
releases (besides the actions undertaken by the Polish Angling Association, where an opin-
ion expressed by an ichthyologist is often outvoted by the democratic decision made by
the PAA’s members and a popular opinion that the more fish are released, the more will
be caught) is in many cases enforced by the regulations – a competition of offers and re-
sulting fishing plans, which include large amounts of fish stock released to water bodies.
Should it serve to support natural fish populations, fish stocking must not be enforced by
the legal system or the pressure of the general public. We should be reminded of an old
rule concerning the maximum allowable catch as the best way to protect fish resources.
From the point of view of economy, inland fishery production is marginal to catches of
marine fish. Considering the costs and means involved in fish stock releases, they seem to
have no economic grounds and the potential mess they cause in gene pools of fish species
living in Poland is enormous. A possible way to reduce this risk is by amending the regula-
tions on leasing fishing grounds. In view of the current “achievements” in this area, a strat-
egy for managing fish genetic resources in Poland should be elaborated. Unfortunately
the costs of such a strategy might be too high versus the potential effects. The question is
whether we are able to bear such costs?
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64
Species, type, and value of
stocking material released into
Polish lakes from 2001 to 2009
Maciej Mickiewicz, Arkadiusz Wołos
Department of Bioeconomics,
Stanisław Sakowicz Inland Fisheries Institute in Olsztyn
Abstract
Stocking lakes in Poland in the 2001-2009 period was characterized by high species
diversity. A total of 23 species of fish and two of crayfish were released into the lakes. In
terms of the quantity of stocking material, the number of enterprises conducting stocking,
the surface area of lakes stocked, and the variety of types of stocking material released,
eleven fish species were of the greatest importance (Crucian and Prussian carp and bighead
and silver carp were each regarded as one species each). Of the eleven species analyzed in
this article, pike has to be considered to be of the greatest importance to the lake fisher-
ies stocking management in Poland, followed by eel, vendace, and carp, then tench and
pikeperch, followed by Crucian and Prussian carp, common whitefish, and wels catfish, and
finally by bighead and silver carp and grass carp. It should also be noted that lake stocking
in Poland during the 2001-2009 period was conducted with highly varied types of stocking
material of the species analyzed. These ranged from the smallest and youngest material
(larvae and montee eel) to reared juvenile types (summer and fall fry) and older types of
material (spring fry and 2+ fry) to adult fish (commercial-sized fish and spawners).
Introduction
The process of eutrophication is one the more significant factors influencing the
intensity and effectiveness of lake fisheries management. However, those who exploit
lakes have a range of management tools at their disposal that can help to curb the nega-
tive impacts of this process. These include fisheries exploitation, protecting fish popula-
tions, and stocking lakes. Stocking is most significant in lakes that are moderately to
heavily disturbed and threatened with degradation, which is the state that the majority
of Polish lakes are currently in (Bnińska 1996).
The importance of stocking is confirmed by the creation of the working party of the
European Inland Fisheries Advisory Commission / Food and Agriculture Organization of
the United Nations (EIFAC/FAO), which developed the main goals for which stocking is
and can be applied (Bnińska 1994). They are as follows:
65
• Mitigation (to compensate for losses in populations caused by environmental changes);

• Maintenance (to maintain fish catches or populations at stable levels when disad-
vantageous changes are caused by the state of the environment);
• Enhancement (to increase catches of a given species);
• Put-and-take (for conducting this type of management for both commercial and
recreational fisheries);
• Put-grow-and-take (as above, and also for both commercial and recreational fisheries);
• Repopulation (for restoring species that have disappeared from environments be-
cause of disadvantageous changes);
• Increasing biodiversity of ecosystems;
• Supplementation (exploiting existing, free ecological niches);
• Providing a feed base for other species;
• Controlling less valuable fish species (populations of fish growing excessively
under the impact of disadvantageous environmental changes);
• Improving environmental quality (primarily through the impact of stocked fish
on other species);
• Protecting ecosystems (understood as maintaining equilibrium among fish popu-
lations).
As this list indicates, the aims of stocking are not strictly focused on management
strategies to obtain defined economic profits from the sale of catches, but they also
include the biological goals of maintaining or increasing aquatic ecosystem biodiversity,
which is exceedingly difficult to assess economically although attempts to do so have
been undertaken (Leakey and Lewin 1999).
Lake stocking management, similarly to any other types of management, must be con-
ducted following the principles of sustainable development, which is also known as ecode-
velopment. In order to achieve this, stocking management must meet three fundamental
conditions: it must be ecologically permissible (safe) while also being socially desirable and
economically feasible (Leopold and Bnińska 1992, Turkowski 2006). One example is the
stocking of lakes with pike. This produces a variety of results, including the increased com-
mercial production of this species, the stimulation of recreational lake fisheries since pike is
the most desirable species among recreational lake fishers (Wołos 1992), and positive regu-
lation of ichthyofauna through increased populations of predators (Leopold et al. 1998).
In Poland in the early 1990s, in addition to such factors impacting lake stocking man-
agement such as the worsening state of lake ecosystems from eutrophication and the
necessity of managing lakes in accordance with the principles of ecodevelopment, an-
other important factor appeared. Namely, this was the ownership transformation of lake
fisheries, which forced fisheries enterprises to function within a free market economy.
This was particularly apparent in eel lake stocking (Leopold and Wołos 2001, Mickiewicz
and Wołos 2002), which, among other aspects of this fishery, radically changed lake
stocking management in terms of intensity (Mickiewicz 2001a, 2006b).
The ownership transformation in lake fisheries resulted in the leasing of so-called
fisheries facilities comprising, among other things, lakes (or even single lakes), and most
of these enterprises were new and included various forms of ownership, such as com-
66
Species, type, and value of stocking material released into Polish lakes…
panies, physical persons, and the Polish Anglers Association. These enterprises were re-
quired by contract to dedicate a designated share of earned income from the sales of
fish catches to stocking the lakes exploited. In the case of contracts with Regional Water
Management Boards, enterprises were required to conduct stocking with the appropri-
ate stocking material of values that were stipulated contractually, which forced enter-
prises to implement and conduct appropriate stocking policies.
Following the fisheries ownership transformation, as noted previously, changes were
made to the lake stocking policy model (Wołos and Mickiewicz 1997, Mickiewicz 2001a,
b, 2006a, b). These were linked to the necessity of adapting fisheries enterprises to the
realities of the market economy and to the increased economic importance of recrea-
tional fisheries (Wołos 2000, 2003), technological advances in the production of stock-
ing material (Mamcarz et al. 1999, Zakęś and Falkowski 2000, Zakęś and Jarmołowicz
2009), as well as to increased ecological awareness, which is reflected in the species
spectrum of lake stocking.
The present article is a brief analysis of the management of lake stocking conducted
in Poland from 2001 to 2009, thus it refers to the period during which the fisheries were
operating within a free market economy following ownership transformation. The anal-
ysis focuses primarily on species biodiversity of stocking, the variety of types of stocking
material of particular species, and the value of the stocking material of particular spe-
cies, which reflects the economic importance of these species in stocking management.
Materials and methods

This article was written based on the following materials:
1. Information regarding the wholesale prices of commercial lake fish and the most
important types of stocking material released by lake fisheries enterprises in 2005
(Mickiewicz 2010).
2. Data from the 2001-2009 period regarding the magnitude of lake stocking col-
lected from annual questionnaires submitted by lake fisheries enterprises exploit-
ing lakes throughout Poland (Tab. 1).
Table 1. Number of enterprises and surface area of lakes exploited according to questionnaires
regarding the magnitude of stocking conducted in the 2001-2009 period
Number of enterprises Combined surface area of lakes exploited

Year
(n) (ha)
2001 65 231 454
2002 65 229 080
2003 66 231 352
2004 64 229 681
2005 95 234 082
2006 91 226 495
2007 104 235 784
2008 129 246 311
2009 104 236 772
67
3. Information from the 2001–2009 period regarding expanded data on lake fisheries
stocking – quantities of stocking material released, surface area of lakes stocked
with particular species, value of stocking material of particular species (Tab. 2). These
data were collected from the same annual questionnaires submitted by lake fisheries
enterprises exploiting lakes throughout Poland as noted in point 2 above.
The analysis was performed with the following methodological approaches and
statistical and mathematical calculation methods.
Table 2. Number of enterprises and surface area of exploited lakes according to complete
questionnaire data regarding stocking conducted in the 2001–2009 period
Number of enterprises Combined surface area of lakes exploited

Year
(n) (ha)
2001 50 149 246
2002 54 192 305
2003 47 169 628
2004 55 204 022
2005 88 220 347
2006 90 224 396
2007 104 235 784
2008 129 246 311
2009 104 236 772
1. The data regarding lake stocking management in the 2001–2009 period was analyzed
statistically with the financial value (pln) of stocking calculated based on the mean
price of particular types of stocking materials used by lake fisheries enterprises in 2005
(Mickiewicz 2010). The financial values obtained were analyzed with regard to various
total surface areas of the lakes studied in subsequent years as the index pln/ha.
2. The only species analyzed were those that were the subject of intense stocking
management in the studied period of 2001–2009. They are:
- eel, Anguilla anguilla (L.);
- common whitefish, Coregonus lavaretus (L.);
- vendace, Coregonus albula (L.);
- pike, Esox lucius L.;
- pikeperch, Sander lucioperca (L.);
- wels catfish, Silurus glanis (L.);
- tench, Tinca tinca (L.);
- Crucian carp, Carassius carassius (L.), and Prussian carp, Carassius auratus gibelio
Bloch., together;
- carp, Cyprinus carpio L.;
- grass carp, Ctenopharyngodon idella Val.;
- bighead carp, Aristichthys nobilis Rich., and silver carp, Hypophthalmichthys mo-
litrix Val., together.
The data collected for this article also indicated that other species of fish and cray-
fish were stocked into Polish lakes during the 2001–2009 period under analysis, as
68
follows: European lake trout, Salmo trutta m. lacustris L.; brown trout, Salmo trutta
m. fario L.; rainbow trout, Oncorhynchus mykiss Richardson; peled, Coregonus
peled Gmel.; perch, Perca fluviatilis L.; burbot, Lota lota L.; asp, Aspius aspius (L.);
ide, Leuciscus idus (L.); roach, Rutilus rutilus (L.); common bream, Abramis brama
(L.); noble crayfish, Astacus astacus L.; narrow-clawed crayfish, Astacus leptodac-
tylus Esch. The stocking of these species was irregular, sporadic, or performed just
once. This does not pertain to the stocking of European lake trout, perch, roach,
or common bream. Although these species were stocked annually throughout the
period analyzed, stocking was done by just a few enterprises at a low magnitude
that was incomparable to values of the stocking of the other species analyzed.
3. The method used for the calculations was time series analysis (trends) taking into
consideration third degree polynomials and exponential curves with a boundary
degree of probability of p ≤ 0.05. Statistically significant polynomials (when there
was a lack of statistical significance in the actual data) are illustrated graphically;
this refers to selected stocking indexes of pln/ha of the species noted above. In the
time series analysis the independent variable was the subsequent number of years
analyzed, and in the case of the 2001-2009 period it was from 1 to 9. The mean
annual value of the stocking species structure in the 2001-2009 period was calcu-
lated, and the results are presented in graphic form.
4. In order to study the differences in stocking management intensity, the following
indexes were calculated for the particular species in the 2001-2009 period:
- share of enterprises (%) performing stocking (number of lake fisheries enterprises
performing stocking of particular species in relation to the total number accord-
ing to data in Table 1);
- share of surface area stocked (%) (the surface area stocked with particular species
in relation to the total surface area of lakes analyzed according to data in Table 2);
- value of stocking with a particular species by lake surface area – pln/ha (for par-
ticular species that were stocked according to data in Table 2);
- value of stocking by the total lake surface area analyzed – pln/ha (for particular
species that were stocked according to data in Table 1);
Mean annual values for the 2001-2009 period were calculated for all of the pre-
ceding indexes, and their basic statistical measures were characterized – standard
deviation (SD) and coefficient of variation (V%).
5. All of the questionnaires that were used in the analysis were conducted according
to principles applied in market research (Sagan 1992, Kotler 1994, Kędzior and
Karcz 1997) and in consideration of the complexity and variation of the problem
under analysis.
Results and discussion

The course of changes in the value of stocking in the 2001-2009 period were calcu-
lated based on questionnaire data (Tab. 1), and on the mean prices of various types of
stocking material utilized in lake fisheries in 2005 (Mickiewicz 2010). Trends were cal-
culated for eleven species of fish that were stocked into exploited lakes by the fisheries
enterprises analyzed: these were eel, common whitefish, vendace, pikeperch, pike, wels
69
catfish, tench, crucian and Prussian carp, carp, grass carp, and bighead and silver carp
(Figs. 1-11). In the cases of common whitefish (Fig. 3), pike (Fig. 4), tench (Fig. 7), and
Crucian and Prussian carp (Fig. 8), only actual data are presented graphically since the
courses of these trends were not significant statistically.
Changes in the values of stocking eel (Fig. 1) were irregular; from the beginning of the
studied period until 2003 they decreased, then increased to the maximum in 2007, only to
decrease again until the end of the studied period. The maximum value of actual eel stock-
ing was noted in 2008 (9.13 pln/ha), while the minimum was in 2003 (5.18 pln/ha).
In the cases of vendace (Fig. 2), pike (Fig. 5) and wels catfish (Fig. 6), trends in the
values of the material stocked decreased initially and then increased. The value of vendace
stocking decreased from the beginning of the period studied to the minimum in 2003, but
then it increased until the end of the period studied, and the maximum was noted in 2009
(Fig. 2). The value of actual vendace stocking was at maximum levels in 2009 (6.82 pln/ha),
while the minimum was noted in 2004 (3.89 pln/ha). The value of stocking pikeperch de-
creased from the beginning of the period analyzed to the minimum in 2005, but then it
increased until the end of the analyzed period to the maximum in 2009 (Fig. 5). The actual
value of pikeperch stocking was at the maximum in 2009 (2.84 pln/ha), while the minimum
was in 2004 (1.44 pln/ha). The value of stocking wels catfish decreased from the beginning
of the analyzed period to the minimum in 2004, then it increased until the end of this pe-
riod to the maximum in 2009 (Fig. 6). The actual value of stocking wels catfish was at the
maximum in 2009 (0.80 pln/ha), while the minimum was in 2004 (0.27 pln/ha).
There was an overall decreasing trend in the changes of the value the stocking mate-
rial of carp (Fig. 9), bighead and silver carp (Fig. 10), and grass carp (Fig. 11). The trend
in changes of the value of carp stocking (Fig. 9) first decreased (until 2002), and then
increased (to the maximum in 2006), and then decreased dramatically to the minimum
in 2009. The actual value of carp stocking was at the maximum in 2007 (6.91 pln/ha),
while it was it the minimum in 2009 (3.92 pln/ha). The trend of changes in the value of
bighead and silver carp (Fig. 10) was irregular; from the maximum in 2001 it exhibited
a decided decreasing trend until 2004, then it increased to the maximum in 2007,
only to decrease dramatically in 2009 . The actual value of bighead and silver carp
stocking was at the maximum in 2001 (0.47 pln/ha), while the minimum was noted
in 2005 (0.03 pln/ha). The trend of changes in the value of stocking grass carp (Fig.
11) decreased distinctly throughout the 2001-2009 study period. The actual value of
stocking grass carp was at the maximum in 2004 (0.25 pln/ha), while the minimum
was noted in 2008 (0.03 pln/ha).
The mean annual value of stocking by species during the analyzed period of 2001-
2009 (Fig. 12) was dominated by pike with a share of nearly 28% of the overall value of
stocking. This species was followed by eel (about 20%), carp (about 15%), and vendace
(about 15%). The value of the stocking of tench, pikeperch, Crucian and Prussian carp,
common whitefish, wels catfish, bighead and silver carp, and grass carp did not exceed
7% of the overall value of stocking for any of these species, while the combined total
for the stocking value of these species was about 23%, of which the value of stocking
bighead and silver carp together and grass carp did not exceed 1% of the total value of
stocking (Fig. 12). The overall mean annual value of stocking the species analyzed in the
70
Figure 1. Changes in the values of eel stocking Figure 2. Changes in the values of vendace stocking
in Polish lakes during the 2001-2009 period. in Polish lakes during the 2001-2009 period.
Figure 3. Changes in the values of common whitefish Figure 4. Changes in the values of pike stocking
stocking in Polish lakes during the 2001-2009 period. in Polish lakes during the 2001-2009 period.
Figure 5. Changes in the values of pikeperch stocking Figure 6. Changes in the values of wels catish
in Polish lakes during the 2001-2009 period. stocking in Polish lakes during the 2001-2009 period.
2001-2009 period was 35.48 pln/ha. The value of stocking these same species during the
1995-2007 period was higher at 40.06 pln/ha (Mickiewicz 2010).
The species structure of the mean values of annual stocking in 2001-2009 period can
be compared to that in the 1995-2007 period (Mickiewicz 2010). It was dominated by
eel with a nearly 31% share of the overall value of stocking. This species was followed
by pike (about 20%), vendace (about 14%), and carp (about 14%). The stocking value
of the other species stocked (tench, common whitefish, pikeperch, Crucian and Prussian
carp, bighead and silver carp, wels catfish, grass carp) did not exceed in any instance 5%
of the total value of stocking, and together was 21% of the value of stocking (op. cit.).
Generally, the only significant change in the species structure was that the dominant
species in the 1995-2007 period was eel and in the 2001-2009 period it was pike.
71
Figure 7. Changes in the values of tench stocking in Figure 8. Changes in the values of crucian
Polish lakes during the 2001-2009 period. and Prussian carp stocking in Polish lakes during
the 2001-2009 period.
Figure 9. Changes in the values of common carp Figure 10. Changes in the values of bighead carp
stocking in Polish lakes during the 2001-2009 period. stocking in Polish lakes during the 2001-2009 period.
During the 2001-2009 period, fisher-

ies enterprises released into lakes various
types of stocking material (different ages)
including larvae, summer fry, fall fry, spring
fry (1+), fry (2+), commercial-sized fish
(consumption size), and spawners, while
the type of eel material were released was
either montee or other reared material
(Tab. 3). In terms of the variety of stock-
Figure 11. Changes in the values of grass carp stocking
in Polish lakes during the 2001-2009 period. ing material type, pike and eel (full range
of stocking material types) and pikeperch
(six stocking material types) dominated. These species
were followed by carp and wels catfish (five types), com-
mon whitefish and tench (four types), Crucian and Prussian
carp, grass carp, and bighead and silver carp (three types),
and vendace (two types). The data presented in Table 3
indicates that lake stocking management in Poland in the
2001-2009 period applied a wide variety of stocking mate-
rial in terms of age and size of the particular fish species
analyzed. This is evidence that the hatchery and rearing fa-
cilities for stocking material are relatively well-developed in
Figure 12. The mean annual value Poland (Zakęś and Falkowski 2000, Zakęś and Jarmołowicz
of stocking by species during the 2009), while also indicating that many of the aims of lake
analyzed period of 2001-2009
(100% = 35.48 pln/ha).
stocking management are being met (Bnińska 1994).
72
Table 3. Types of stocking material (X) of particular fish species released

into Polish lakes in the 2001-2009 period
summer commercial
species larvae fall fry 1+ fry 2+ fry spawners
fry fish
vendace X X
common
X X X X
whitefish
pike X X X X X X X
pikeperch X X X X X X
wels catfish X X X X X
tench X X X X
Crucian
X X X
and Prussian carp
carp X X X X X
grass carp X X X
bighead
X X X
and silver carp
montee1) other stocking material2)
eel X X
1)
eel glass fry (montee) – more than 1500 individuals per 1 kg of fry
2)
eel stocking material other than glass fry
Table 4 presents the mean annual values of various coefficients that characterize lake
stocking management for particular species in the 2001-2009 period.
In terms of the share (%) of enterprises conducting stocking, the index was the high-
est for pike, and its stability over the years analyzed was also high. More than half of the
enterprises analyzed also stocked carp, tench, and eel, while slightly fewer than 50% of
the enterprises stocked pikeperch and vendace; however, in these instances the stability
of stocking was not as high. This was also noted with regard to Crucian and Prussian
carp, which was stocked by more than 30% of the enterprises analyzed.
In the case of the share (%) of surface area stocked, pike also dominated, followed by
eel, tench, pikeperch, and vendace. The standard deviation and coefficient of variation
for this index did not indicate such stability during the analyzed period as they indicated
for the previous index.
The next index presented in Table 4, which is the value of stocking by surface area of
particular species (pln/ha), was the highest with regard to carp, but, again, the values of
the standard deviation and the coefficient of variation were relatively high. The subse-
quent species that were noteworthy are vendace, common whitefish, pike, and eel, for
which the coefficient of variation for the value of the stocking of these species did not,
in any instance, exceed 40%.
The final index presented in Table 4 is the value of stocking for total surface area
analyzed (pln/ha). The values of the stocking of pike, eel, carp, and vendace dominated
in this instance, and the standard deviation and coefficient of variance indicated strong
stability in the stocking of these species throughout the total surface area analyzed.
73
The same indexes presented in Table 4 were analyzed for lake fisheries management
for the 2001-2007 period in Mickiewicz (2010). The comparison of these indexes with
those from the 2001-2009 period indicated that there were no significant differences or
distinct changes.
Table 4. Mean annual values of selected coefficients characterizing lake stocking management
of particular species in the 2001-2009 with statistical measures of these values
– standard deviations and coefficients of variation
Coefficient/statisti-
Species
cal measure
grass carp
pikeperch
and silver
whitefish
common
Prussian
vendace
bighead
Crucian
catfish
tench
wels
carp
carp
carp
pike
and
eel
share of enterpris-
50.56 47.11 31.44 86.56 48.44 25.56 55.89 36.33 51.67 10.11 7.44
es stocking (%)
standard deviation
6.71 8.46 8.26 3.50 4.95 4.48 6.88 4.87 8.26 4.17 4.13
(SD)
coefficient of
13.27 17.96 26.28 4.05 10.22 17.51 12.31 13.41 15.99 41.21 55.43
variation (V%)
share of surface
58.37 19.43 15.40 70.42 21.47 8.21 26.40 11.84 14.31 1.09 0.97
area stocked (%)
standard deviation
10.22 2.33 3.33 7.13 6.66 3.52 6.33 3.25 3.73 0.66 0.62
(SD)
coefficient of
17.50 11.97 21.60 10.13 31.04 42.91 23.98 27.45 26.08 60.30 63.98
variation (V%)
value of stocking
by surface area of
12.04 24.37 14.98 13.90 10.63 7.61 7.39 8.80 35.26 10.46 16.96
particular species
(pln/ha)
standard deviation
2.20 4.04 5.92 1.04 3.54 1.84 1.84 1.86 12.55 5.14 14.69
(SD)
coefficient of
18.30 16.59 39.54 7.49 33.28 24.21 24.86 21.18 35.60 49.17 86.62
variation (V%)
value of stocking
for total surface
6.91 5.22 1.26 9.85 2.18 0.53 2.37 1.42 5.44 0.14 0.16
area analyzed
(pln/ha)
standard deviation
1.30 1.06 0.46 1.77 0.46 0.16 0.70 0.44 0.87 0.09 0.13
(SD)
coefficient of
18.73 20.25 36.92 18.02 21.31 30.90 29.72 30.74 15.95 67.89 81.53
variation (V%)
Conclusions
To summarize briefly the preceding considerations, it can be concluded that stocking
lakes in Poland in the 2001-2009 period was characterized by high species diversity. A total
of 23 species of fish and two of crayfish were released into the lakes. In terms of the quan-
74
tity of stocking material, the number of enterprises conducting stocking, the surface area
of lakes stocked, and the variety of types of stocking material released, eleven fish species
were of the greatest importance (Crucian and Prussian carp and bighead and silver carp
were each regarded as one species each). Of the eleven species analyzed in this article,
pike has to be considered to be of the greatest importance to the lake fisheries stocking
management in Poland, followed by eel, vendace, and carp, then tench and pikeperch,
followed by Crucian and Prussian carp, common whitefish, and wels catfish, and finally by
bighead and silver carp and grass carp. It should also be noted that lake stocking in Poland
during the 2001-2009 period was conducted with highly varied types of stocking mate-
rial of the species analyzed. These ranged from the smallest and youngest material (larvae
and montee eel) to reared juvenile types (summer and fall fry) and older types of material
(spring fry and 2+ fry) to adult fish (commercial-sized fish and spawners).
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i nadbrzeżnych obszarów rybackich w województwie warmińsko-mazurskim [Review of the current
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the Warmian-Mazurian district] [Ed. Wołos A.], Wyd. IRS Olsztyn: 95-110. (in Polish).
76
European grayling (Thymallus
thymallus L.) stocking material
production in the context of
the conservation of natural
biodiversity of the environment
and active protection of the
species.
Mariusz Szmyt
Department of Fish Biology and Breeding, University of Warmia

and Mazury, ul. Oczapowskiego 5, 10-719 Olsztyn,
e-mail: mariusz.szmyt@uwm.edu.pl
Abstract
The paper contains information on the biology, ecology and current status of the Eu-
ropean grayling (Thymallus thymallus) in Polish waters. Based on the available literature,
the distribution of the species in Europe and Poland has been described alongside its
environmental preferences, threats in the wild and future prospects. Possible ways have
been suggested of relieving the tension and compensating the species for the unfavour-
able changes in the environment. Guidelines for the active protection of grayling have
been discussed as well as their practical applicability and potential problems associated
with their implementation. The importance of grayling stocking material production
based on farmed broodstock has been indicated. Finally, the role of people in re-crea-
tion and sustenance of the natural environment biodiversity, of which the grayling is an
inherent element, has been reviewed.
Introduction
The European grayling (Thymallus thymallus L. 1758) belong to the salmonids (Sal-
monidae), the subfamily of Thymallinae and the genus Thymallus. Grayling inhabits ex-
clusively the freshwaters of the northern hemisphere, mainly brooks and rivers, mountain
or submountain in character, as well as some clean, ultra-oligotrophic lakes, e.g. Lake
77
Baikal, Kozgol or Imandra

(Witkowski et al. 1984).
The distribution of
grayling in Europe is il-
lustrated in Figure 1. The
southern boundary of the
occurrence of this species
reaches the northern parts
of Italy, Macedonia and
Bulgaria. To the west, it can
be found as far as France
and the UK, whereas to the
east its range of occurrence
ends by the Ural Moun-
tains. Northwards, grayling Figure 1. Range of natural occurrence of grayling in Europe
(Lusk et al. 1987).
spreads nearly all over the
Scandinavian Peninsula.
Nowadays, the pres-
ence of grayling in Poland
is mainly limited to mor-
phologically highly diverse
watercourses, aggregated
in two large regions – the
north and the south of the
country (Fig. 2).
Apart from the natu-
ral habitats, grayling has
been introduced to water-
courses in which it did not
appear before. Particularly
successful introductions of
this species have been per-
formed to the rivers Sanok,
Wisłok and Ropa. In these
Figure 2. The current distribution of autochthonous (filled circles) rivers, below dams, gray-
and introduced (empty circles) populations of grayling in Poland ling has found excellent
(Augustyn and Cieśla 2000)
conditions. The regulated
flow and stable thermal conditions enabled the fish to form strong and naturally re-
producing populations.
Positive effects have also been obtained with regard to introductions of greying to
the Pasłęka (Kostecki 2000), the Tanew, Wisłoka, Dłubnia, Rudawa, upper Pilica, Warta
78
European grayling (Thymallus thymallus L.) stocking material production…
and Vistula, in the Wełna, Drwęca, Supraśl and Bystrzyca (a tributary of the Wieprz River)
(Augustyn and Cieśla 2000).
Environmental preferences, current status and threats

The grayling is a typical rheophilc fish (Mallet et al. 2000). It shows distinct environ-
mental preferences depending on the developmental stage (Sagnes et al. 1997). There
is a very clear relationship between the morphological features of that fish and the
physical properties of the environment it dwells in. In general, as the fish grows, juvenile
forms of grayling migrate from off-shore, shallow sites, with a slow water flow and fine
substrate covering the bottom, to deeper sites, closer to the main current, with a more
rapid water flow and a harsh-grain substrate (Bardonnet et al. 1001, Sempeski and Gau-
din 1995). It has also been discovered that juvenile grayling change sites daily, between
foraging grounds and benthic resting areas, very shallow ones and occupied at night
(Gaudin and Sempeski 2001).
Older grayling (fish fry in the second year of life and adult individuals), depending on the
size of a watercourse and season of the year, prefer relatively deep sites, with a fast flow of
water and larger grain size of the substrate (Vehanen et al. 2003, Nykänen et al. 2004).
Grayling is a short-lived fish. Within our geographical latitudes, specimens more than
5 years of age make up a small percentage of populations. Reports published by many
authors (Woolland and Jones 1975, Witkowski et al. 1989) suggest that most of the fish
obtained for their studies were no more than four years old (3+). Grayling over 5 years
of age were 4% at the most of all captured fish.
Many factors affect the species composition of the ichthyofauna inhabiting a given wa-
tercourse (Wiśniewolski 2002). With respect of grayling, the main factors are: quality of
water (temperature, oxygen concentration, chemical composition), depth of a watercourse,
rate of water flow and presence of natural depressions and obstacles where fish can hide.
The optimum range of temperatures for grayling is from 4 to 18oC (Elliot 1981, after
Crisp 1996). Temperatures below or above the optimum are claimed by that author to
be critical for grayling. As Ring (1956) stated, grayling is more sensitive than trout to
excessive warming up of water and dies when the water temperature rises above 20oC.
But to be more precise, Harsány and Aschenbrenner (2002) suggest that grayling lose
their adaptability to higher temperatures in the second year of age, while during the
first one they tolerate temperatures above 20oC. These observations, however, are con-
tradicted by the fact that grayling are present in such rivers in Pomerania (Pomorze) as
the Radunia, Brda or Drawa, where summer temperatures of water are often above 21oC
(Goryczko K., Inland Fisheries Institute in Olsztyn, Department of Salmonid Research in
Rutki, oral report).
The water purity status, and especially presence of any industrial contaminants or
toxic effects of various chemical elements and compounds, is another fundamental as-
pect which determines whether grayling can survive (Vuorinen et al. 1998, Honkanen et
al. 2005). Apart from pollution, other examples of transformations of the natural envi-
ronment caused by man can be indicated as having an adverse influence of ichthyofauna
(Penczak and Kruk 2000, Wiśniewolski 2002), for example constructions that cut a river’s
continuity or regulations of river channels. These can make migrations of fish impossible
79
and prevent them from reaching their spawning grounds. An example to confirm it is
the construction of a weir on the Leśnica River (a right tributary of the Dunajec River)
– about 600 m from its outlet. The height of the weir up to about 0.5 m made it utterly
impossible for grayling to continue their annual spawning migrations upstream the river
(Kowalewski M. the PAA, region in Łopuszna, oral report). Effects of building construc-
tions across river channels have also been described by Ovidio and Philippart (2002).
These authors, however, added that factors like the design of a construction, water
temperature and its depth below the barrier all have an impact on how effectively the
fish can overcome that obstacle.
Many mountain and submountain rivers have changed their character completely
due to human actions. Thermal conditions in these rivers have deteriorated drastically, as
the range of temperatures and rates of their changes have increased (Goryczko 2003). In
summer, the water temperature in the ‘civilized’ sections of the Raba, Czarny Dunajec or
Kamienica Nawojska may range from 10-12oC in the morning to 24-26oC in the evening.
In winter, the bottom in shallow and overflowing rivers may freeze and pancake ice
forms more readily.
Another example of degradation of an aquatic environment is the mining of boul-
ders, gravel and aggregate from river channels (Ring 1956, Goryczko 20030. This is espe-
cially noxious due to the damage caused to spawning grounds and sites where juvenile
individuals grow, and it affects other fish species apart from grayling. For a watercourse
to offer suitable conditions for salmonid fish, it needs to contain quite a broad grain size
spectrum of gravel, as the role of a substrate changes while the fish grow and develop
new requirements (Kondolf 2000).
A huge impact on the abundance of grayling populations is produced by the inci-
dence of natural disasters, which in a sense are a consequence of human activities. In
July 1997, the territories of the Czech Republic and Poland were struck by the most cata-
strophic flood during the 20th century. In the watercourses directly exposed to the flood
wave, a considerable decrease in the abundance of the populations of grayling was
observed, especially among older fish (Lojkásek et al. 2005). During the two years after
the flood, the recorded angling catches comprised significantly fewer larger specimens
than before the disaster. As Augustyn (2002) claims, ‘the century’s flood’ could have
annihilated the whole fish fry of that year, and another flood, in the late April and early
May of 1998, although much smaller, could have destroyed the laid spawn. Prokeš and
Baruš (1998), who investigated the state of the ichthyofauna in the Morava River im-
mediately after the flood, discovered elevated numbers of grayling in the lower sections
of the river. But the authors suggested that the migration of fish downwards the longi-
tudinal profile of the river may have been forced by the receding flood wave. A similar
phenomenon was observed by Lojkásek et al. (2005). Moreover, the increased number of
fish individuals was accompanied by a decrease in the biomass of fish. This was caused
by the fact that masses of water swept individuals belonging to younger age groups.
Another important, albeit frequently overlooked, factor influencing the abundance
of grayling is the predatory pressure of fish-eating birds. Spurný and Guziur (2002) re-
port that cormorant pressure had reduced the population of grayling in the Beczva River
in the Czech Republic by nearly 60%. Grayling from the Alz River, in southern Germany,
80
were also found to be an important component of the cormorant’s diet (Keller 1998).
Other bird species which may exert strong predatory pressure on grayling are storks and
herons (Kowalewski M., the PAA region in Łopuszna, oral report).
Among several, identified reasons why the number of grayling in Poland has been
limited, the key role seems to be played by a growing angling pressure and poach-
ing (Augustyn 2002, Holčik 2004). These two researchers point mainly to the dynamic
growth in the number of anglers as well as to the continually improved angling tools
and techniques. It seems that in some social circles there is tacit consent to illegal and
wasteful fish catching. Fish poaching brings more harm because in many cases the gray-
ling caught are young individuals before first spawning.
Species protection and prospects

The aggregate effect of all the factors discussed in the previous section of this article
makes the future existence of grayling in Polish waters depend on all kids of protection
treatments. Grayling in the Odra River basin is classified as a vulnerable (V) species,
which means it faces a high risk of becoming extinct in not so distant future. In the Wisła
River basin, the species is near threatened (NT) and belongs to the category of lower risk
(LR) (Witkowski et al. 2004). The species is also listed on the Red List of Fish and Lam-
preys of 2009 (Witkowski et al. 2009). Moreover, the question of grayling protection has
drawn attention of persons and institutions drafting and passing many legal documents
and acts. The Bern Convention of 1979, ratified by Poland in 1996, or the Ordinance of
the Minister for Agriculture and Rural Development of 2001, on fish catches, fish culture
and rearing and catches of other aquatic organisms (Journal of Law 1996, JoL 2001) are
examples of such regulations.
One of the ways of counteracting the diminishing abundance of grayling in open
waters is the so-called “active protection of a species”. Its basic component is ex situ
protection, which in the case of fish means mastering the methods for breeding a given
species and maintaining its farmed broodstock in fish ponds (Goryczko et al. 2001).
An important component of “active protection” is semen cryopreservation, successfully
worked out and implemented for grayling (Lahnsteiner et al. 1996).
Another, successful way to protect grayling is by reconstructing degraded spawning
grounds (Zeh and Dönni 1994). The benefits derived from this approach are double. We
not only recreate potential spawning grounds but also provide juvenile grayling originat-
ing from fish fry releases with suitable places to dwell.
Considering the complexity of the problem of active protection and the fact that it
may be solved with an aid of a combination of many disciplines, it seems important that
activities carried out on different levels should be interconnected. Logically, however,
with the “key factor” mentioned earlier, we should pay closer attention to supporting
natural populations with the material obtained under controlled conditions.
Fish stocking generate different costs. Financial costs are easily calculable, but envi-
ronmental ones are much more difficult to forecast, and the results of our fish stocking
treatments can surface after many years. Unfortunately, they can have highly negative
impact. Human interference in a relatively stable system of local, natural populations of
grayling by introducing reared fish fry may disturb distinct gene pools of these popula-
81
tions (Koskinen et al. 2001, Jankun et al. 2003). This holds true for other endangered fish
species as well, all of which are in inherent component of the biodiversity of the natural
environment. Fish fry releases not based on rational and scientific grounds, instead of
yielding expected results, may bring damage to the fitness and stability of supported
populations (Łuczyński and Bartel 1997).
It is unacceptable to release to open waters fish stocking material of uncertain origin.
Uncontrolled fish stocking, often with the material purchased in the Czech Republic or
Slovakia (Bartel 2004), destroys biodiversity and does more harm than benefit.
However, man’s activity in terms of supporting natural populations with material ob-
tained under controlled conditions does not mean just causing losses and creating threats.
Fish breeders’ experience, higher ecological awareness and constantly improved methods
of grayling breeding enable us to look more hopefully into the future. The complex ap-
proach to the problem, i.e. from the moment spawn is obtained and incubated, through
the rearing up of fish fry until grayling selects and broodfish are produced, gives us
a chance to compensate for the losses caused by human actions (Kowalewski 1987, Grud-
niewska and Szmyt 2003, Szmyt et al. 2004). Furthermore, since it is possible to verify the
effectiveness of fish fry releases by tagging juvenile stages of grayling (Nagięć et al. 1995),
we have been able to create the fish stocking policy in a rational manner and in accord
with the specific environmental requirements of the species (Goryczko et al. 2004).
According to the data of 2000 (Augustyn et al. 2000), stocking material of grayling
in Poland was produced in 8 trout farms. In contrast, there were 28 such trout farms in
the Czech Republic (Pokorný 2000). Nonetheless, in both cases, the authors imply that
there has been a regress in grayling stocking material production. A possible reason
could be the difficulties encountered in developing the grayling production technology.
Many farms continue to obtain wild broodfish from the natural environment. However,
in a situation where the abundance of populations of this species has been drastically
reduced, such actions are less effective and can make natural spawning more difficult.
Farmed broodstock
A possible and successful solution to the above problem is maintaining farmed brood-
stock under controlled conditions. Typical trout farms handle it very well (Witkowski
and Bartel 1999), although the biotechnique of farming grayling, compared to rainbow
trout, poses many more problems (Kowalewski 1987, Goryczko et al. 1995, Szmyt et
al. 2004). A grayling farmed broodstock is the essential element of all stocking pro-
grammes, included in species active protection initiatives. Therefore, reproducing fish
should be provided foraging and living conditions as closely similar to the natural ones
as possible. Despite all efforts, keeping grayling fish for reproduction under controlled
conditions is associated with considerable losses, up to 50% of the total number of
broodfish (Harsány and Aschenbrenner 2002, Szmyt and Grudniewska 2005).
One of the major problems faced in rearing grayling fish for reproduction in fish ponds
is the fish loss after spawning. The actual loss, however, varies from year to year and
depends on many factors. Mortality of reproducing fish can be greatly reduced if we are
extremely careful while manipulating fish, apply anaesthesia before obtaining gametes
and give fish disinfectant and medicinal baths afterwards. According to Kupka (1967) and
82
Kowalewski (the PAA Region in Łopuszna, oral report), post-spawning loss in a maternal
breeding herd of grayling should not exceed 5-10%. But when extremely negative thermal
and health conditions appear, such loss can reach up to 60% (Grudniewska and Szmyt
2003). With high losses, it is difficult to build a stable farmed broodstock comprising fish
of all ages. A higher mortality among older farmed broodfish leads to a situation when
up to 80% of obtained eggs come from two-year-old fish (Grudniewska et al. 2001). Loss
among farmed broodfish is also due to the need to check on their sexual maturity. There-
fore, during reproduction, it is necessary to repeat manipulations with brodfish many
times. And as they are sensitive to all kinds of skin abrasions and damage, secondary bac-
terial or fungal infections are likely, and these lead to fish loss.
This problem can be solved through application of hormonal stimulation of repro-
duction (Szmyt et al. 2007). Because of the anatomical structure of females of salmonid
fish, it is easy to obtain their gametes. However, when spawning is stretched in time, it
causes problems. Should we apply hormonal stimulants, we could induce ovulation and
synchronize spawning (Arabaci et al. 2004). Being able to achieve such an effect would
be another step forward in mastering the biotechnique of breeding this species under
controlled conditions.
One of the essential aspects of maintaining farmed broodstock under controlled con-
ditions is the question of feeding fish properly and strictly according to their needs. With
regard to species which do not have commercial value as food resource, including gray-
ling, it is difficult to hope that full value feed for their broodfish will be manufactured.
Therefore, in order to achieve the set aim, feeds for brodfish should be supplemented
so as to compensate for deficits of some nutrients. Supplementation, however, means
elaboration and implementation of a suitable technology, which will require financial
outlays. The latter aspect is of particular importance in the case of non-commercial fish
species. Addition of nutrients supplementing fish diet should be economically accept-
able and have positive influence on production effects (Szmyt et al. 2004).
In many cases, efforts undertaken to optimize nutrition of farmed broodfish are not
satisfactory. It is not easy to imagine how, under controlled conditions, we can provide
fish with optimum nutrition. Nevertheless, construction of artificial river channels as well
as keeping fish in supplying channels and settlement tanks, gives us such an opportu-
nity. Conditions being similar to natural ones can satisfy the needs of broodfish most
completely, both in terms of living and feeding requirements. This, obviously, means
their better shape and better biological value of gametes.
Optimization of the technology for production of grayling stocking material gives
a chance that the decline in the number of grayling fish in Polish waters will be radically
restrained, provided that the existing legal and administrative norms set up to protect
the species will be respected (Szmyt and Grudniewska 2005).
Moreover, if we strongly object to thoughtless destruction of natural populations of
this species and insist on some amendments to the law regulating its protection (Augustyn
2004), we should be able to accelerate the process of restoration of this beautiful fish to
its natural habitats. Somewhat of a curiosity could be the information cited by Cios (2007)
that back in the Middle Ages grayling used to be a valuable component of rents and pur-
chases made, for example for Queen Jadwiga and King Władysław Jagiełło.
83
Summary
When analyzing the examples described in this paper, it seems justifiable to consider
the question of “active protection” of grayling in many aspects. This species, doubtless
a unique and environmentally valuable one, is an inherent element of natural biodiver-
sity. It is paradoxical that although many negative transformations of the environment
are caused by people, further depletion of the grayling populations in Polish waters and
degradation of natural biodiversity is unavoidable unless people help the species.
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86
Utility and market value of lakes
Konrad Turkowski
Department of Spatial and Environmental Economics / Department

of Lake and River Fisheries, University of Warmia and Mazury in
Olsztyn, kontur@uwm.edu.pl
Abstract
Three basic functions of lakes, which determine their economic values, have been in-
dicated: a resource of water for human uses, a real estate and a natural resource. Within
these three functions, the following water uses (detailed functions) have been distin-
guished: angling and other types of water recreation, fishery, landscape shaping, biodi-
versity, water retention and regulation of water relations, reception and assimilation of
pollutants, etc. The choice of a function which serves to estimate a lake’s value depends on
the purpose and extent of the appraisal and in turn determines which appraisal methods
will be applicable. Lakes as resources of water for human and industry uses are a product
of the volume and quality of water they store as well as of their location. The economic
value of such resources has been presented as a sum of differential rents achieved from
having water resources of the quality, retention and localization better than the margin
ones. The value of a lake as a real estate does not reflect its value as a natural resource,
because an appraisal of a real estate property does not extend beyond the scope of own-
ership rights and consequent benefits derived by the owners. Among the characteristics
which shape the transaction prices of a lake are the lake’s location, quality of water and
trophic state of the lake, the character of the lake’s drainage basin, landscape and recrea-
tional assets as well as the lake’s suitability for fishing and angling. The conditions under
which lakes can enter the real estate market as well as the approaches, methods and
techniques employed for their appraisal have been presented in accord with the relevant
obliging law. The comparative approach to the market valuation of lakes has been illus-
trated with examples from practice.
Introduction
Value is a very broad concept. It can be considered in the economic sense, as much as
in terms of aesthetics, ethics or morality. In economy, valuation generally means estab-
lishing monetary values we are willing to sacrifice in order to own or use given goods or
services. These values depend on the utility and accessibility of these goods and services,
which, in the market economy and in accordance with the rules of supply and demand,
are reflected as transaction prices.
87
In an appraisal of things, events or actions, certain values are assigned according to

a scale proper for a given object and scope of the valuation, e.g. valuation of aesthetic
assets requires that a scale from beauty and ugliness be taken. It raises no doubt that
such evaluation is largely subjective and depends on who makes it. It is even more diffi-
cult to evaluate universal values like respect, honour or honesty. Similar problems are en-
countered when natural assets are to be evaluated, because then altruism, for example,
is a specific value created on the basis of people’s awareness that a given natural asset
should be preserved (McConnel 1997, Johansson-Stenman 1998, Wiekard 2005).
The utility value of lakes
The utility value defines how given goods can satisfy human or, in a broader context,
environmental needs. The use value exists independently of the availability of a given
object on the market or individual preferences of market participants. Some goods can
have a very high use value but a relatively low market value due to their availability on the
market (e.g. air or water) and, reversely, when in shortage on the market, some goods can
gain high market values even though their use values are low. Nevertheless, lack of use val-
ues equals lack of economic values. Obviously, the highest market values will be assigned
to the goods of the highest use value which are rare and difficult to obtain, for example
diamonds. In other words, the use value conditions the market value of a given thing, but
is not subjected to the supply and demand rule, which governs market values.
Valuation is an action or a series of procedures whose aim is the determine value.
However, it is extremely difficult to establish one, absolute value for a given resource. As
a rule, the economic value is relative. It depends on the scope of valuation and on the
conditions under which it is conducted. In practical measures, the value is a derivative of
the function or usability of a given resource.
Table 1. Scope of a lake appraisal depending on the lake’s functions and uses.
Basic functions of lakes

Detailed functions/uses (elements of appraisal) Resource of water for Nature
Real estate
human uses resource
Fishing/angling no* yes yes
Recreation no yes yes
Retention/water storage/regulation
yes no yes
of water relations
Potable water/water for other humans uses yes no/yes yes
Component of landscape no yes yes
Biodiversity no no** yes
Receptio nand decomposition of pollutants *** no no yes
* - under certain conditions, fishery treatments can be performed as an element of water quality protection
(Falkowski 2006).
** - under normal conditions, biodiversity is not a dominant price-shaping feature in valuation of lakes as real estates.
In some specific cases, it can depress the market value of a lake, e.g. when for the sake of nature resources conserva-
tion, the rights of a potential owner are limited, e.g. fishing forbidden in a lake situated in a nature reserve.
*** - the use of a lake as a reservoir to receive and decompose pollutants when it functions as a reservoir of potable
water and water for other human uses and a real estate is neutral as long as it does not cause any deterioration
of the quality of water in the lake. With respect to a lake as a nature resource, this function is positive (the lake
participates in the matter and energy cycle in a given environment) as long as the load of pollutants does not
disrupt the natural ability of the lake for self-purification. For more information on the ability of lakes to assimilate
pollutants and on their vulnerability to anthropological influences, cf. Kudelska et al. (1991).
88
In terms of the utility of lakes, three basic functions can be named: a resource of wa-
ter for human and industry uses, a real estate property and a natural resource (Turkowski
2003). These functions can be assigned a number of detailed functions (uses), which to-
gether determine the economic value of lakes (Tab. 1). Depending on the functions they
perform, lakes can carry different values, which may be manifested through the applica-
tion of various, sometimes completely different methods. Which of the above functions
will lie at a base of valuation depends on the purpose of the process, and that in turn is
determined by the intended use in spatial development, type and location of a lake.
The value of a lake as a resource of water for human uses

The value of a lake as a resource of water for human uses, including potable water
resources, is a product of the volume and quality of water in the lake and the lake’s
location. For human uses, the most valuable are clean waters, of high quality, situated
at a convenient, short distance. When many such lakes are available, their market value
(or derivative economic value) will be low despite their high use values. However, the
resources of clean waters are largely limited and their replenishment incurs high costs
of water treatment or import from other regions. Such outlays would most probably
be reflected in the purchase prices for water. The problem lies in the fact that due to
political and legal as well as social considerations, water resources are not submitted to
free turnover, and prices or rather fees for using water do not reflect the actual costs. In
this context, the economic value is best manifested by the differential rent, which is the
difference between the costs of treating and acquiring the resources possessed and the
costs of managing the margin resources. The margin resources are the ones whose costs
of management are equal to the profits obtained. Acquisition and treatment of waters
worse than margin ones incur losses, which rise proportionately to the deteriorating state
of waters and conditions under which they are obtained. In the water economy, costs
saved by possessing water resources of the quality and location better than the margin
ones and less expensive to retain add up to specific water rents (Turkowski 1994):
W = Wp + Rj + Rl + Rret / 1 /
where:
W – value of water in a given water body in PLN/m3,
Wp – average value of a unit of water in a given region or river basin in PLN/m3,
Rj – rent from water quality in PLN/m3,
Rl – rent from location of a water body in PLN/m3,
Rret – rent from water retention (e.g. in a lake versus the maintenance
and appreciation costs of a dammed reservoir) in PLN/m3.
The sum of the above rents reflects the value of annual “savings”, i.e. the surplus of
the value over the current value of the margin resources assumed to be zero. The value
of water resources assessed as above should be taken into consideration when making
decisions on water management, especially when setting fees or compensations for pol-
luting or degrading water resources. In order to move on to the proper economic valua-
tion of resources, one would have to capitalize all the above annual values. Considering
89
0 0
Location,
good good 50 2922 0
access
Fish management in a variable water environment 100 5345
Nature and 0 0
that they last over tens of years at least, the simple capitalization technique should be
landscape moderate good 50 213 -21
sufficient (Turkowski and Lirski 2010).
values 100 426
Lakes as real estate properties

0 0
Fishing
Lakes are real estate properties, and boundaries are set by register shorelines (The
whose good very good 50 61 -6
Act on Water Law, referred to as the Water
angling values Law, Journal of Law No 239, item 2019 with
100 122
amendments). However, not all lakes can be freely sold and purchased. In the light of the
current law, lakes are classified as inland surface flowing or stagnant Sum waters (Fig. 1). - 27
Flowing water are waters in natural watercourses, channels and in springs from
Source: Turkowski 2009 m
which watercourses originate. In legal terms, lakes are flowing waters if they have con-
stant or periodic inflow or outflow of surface waters or if they are artificial lakes situated
along the course of flow-
ing waters (Fig. 1). In land INLAND SURFACE WATERS
registration ledgers, flow-
ing waters are marked with
the symbol (Wp). Flowing waters Stagnant waters
Stagnant waters are

lakes and other natural
water bodies which are Lakes and other natural
water bodies Lakes
not connected directly, in Rivers
(so-called open) (so-called closed)
a natural way, with surface Other natural
flowing waters. In land Mountain Storage water bodies
streams (so-called closed)
registration ledgers, they reservoir
are marked with the sym-

Figure 1. Legal division of inland surface waters.
bol (Ws). Figure 1. Legal division of inland surface waters.
The division of inland surface waters, including lakes, into flowing and stagnant
ones is of the utmost important to their legal ownership. Waters classified as flowing
ones belong to the State Treasury, are public waters and are excluded from the market
(cannot be sold and purchased). In turn, stagnant waters, including lakes thus classified,
belong to the owners of the land properties within which they are localized. Along with
the grounds, these lakes can be sold and bought on the market.
In the light of the above legal regulations, it is only the lakes classified as stagnant
waters (by convention, non-discharge or closed lakes) that can be an object of market
turnover. Lakes classified as flowing waters belong to the State Treasury and cannot be
submitted to the law on real estate properties.
The market value of lakes

Appraisal of lakes as real estate properties is associated with the valuation of rights
that potential owners or users will acquire, and this, to a large extent, depends on the
relevant legal solutions. Another factor of equal importance with respect to the market
value of lakes is the question of their availability on the market, and mainly – the number
of offers to sell and conditions imposed on the turnover of lakes.
Rules applicable to the appraisal of a real estate – regardless of it type, location or
intended use, and likewise independently of the owner – are regulated by the Act on real
90
estate property management of 21 August 1997 and subsequent executive regulations

(JoL of 2010, No 102, item 651, with amendments). In compliance with these regula-
tions, a market value of a real estate can be determined by using appraisal methods
encompassed by the comparative or the revenue approaches or else a mixed approach, if
it comprises elements of the former two approaches (Tab. 2). The choice of an appraisal
method depends on the scale of an appraisal, type and location of a real estate, the lo-
cal development plan, the present condition of the real estate and available information
about prices of similar real estates in the course of market transactions.
Table 2. Approaches, methods and techniques for appraisal of the market value
of a real estate in compliance with the binding law on real estate management.
Approach Method Technique

1. pair comparison
Comparative 2. correction of average price no specified techniques
3. statistical analysis of the market
1. investment 1. simple capitalization
Revenue
2. profits 2. cash flow discounting
1. residual
Mixed 2. land estimate indices no specified techniques
3. liquidation costs
The market value of lakes in the comparative approach

The comparative approach relies on the determination of the value of a real estate
on the assumption that this value is equal to the prices for similar real estates obtained
in the course of market transactions.
Considering the fact that the turnover of lakes is very small, their valuation is most
often based on comparisons of pairs. Specification of the value of a real estate with this
method involves its comparison, one by one, with at least three similar real estates of
known properties and obtained prices. Other methods are applicable on condition that
a large number of market transactions are known, from dozens of transactions (the
market statistical analysis method) to at least between ten and twenty (the average price
correction method) (Tab. 2). The actual number of closed buy-sell transactions for lakes,
even countrywide, is typically too small to satisfy that condition.
The procedure for valuation with the pair comparison method consists of the follow-
ing steps:
- assessing the legal and user status of a given lake, specifying its basic features
(called attributes) which determine its value,
- searching for lakes as closely similar to the appraised one as possible on the trans-
action market,
- determining the effect of time on levels of prices for lakes in the same area and,
accordingly, correcting the appraised value,
- based on a market analysis or behaviour of buyers, specifying the weights of
basic features (attributes) of lakes which shape their transaction prices on this
market,
91
- making at least three pairs of the appraised lake with the lakes taken for com-
parison and correcting their prices by the value of the total correction amounts,
according to the previously adopted attributes,
- calculating the unit (in PLN/ha) and total (in PLN) value of the appraised lake.
Assessment of the legal status mainly means the determination of the form of owner-
ship, the data on the lake in the land registration system, its intended use in the local spatial
planning or in the land use planning for a given commune (gmina), as well as on the current
condition of the lake. Attention should also be drawn to any legal or administrative limita-
tions imposed on the lake’s use and originating from a possible special use of the water
body, e.g. if the lake lies in a nature protection area or in a potable water intake zone.
Basic attributes of lakes and their valuation

The choice of basic attributes, i.e. the features which most probably have a predomi-
nant influence on the level of transaction prices for lakes, depends on local conditions.
There are, however, features which are above the local importance, shared by most
lakes. Among such most important properties of lakes, which are usually taken into
consideration in the valuation of lakes, are (Turkowski 2003):
- location,
- quality of water and the trophic state of a lake,
- the character of the drainage basin and threats,
- landscape values,
- fishing and angling utility.
Location of one of the most essential attributes of a real estate. General and de-
tailed locations are distinguished. The general location specifies the location of a lake
in a given region, distance to big agglomerations as well as to the nearest villages or
towns. A short distance to urban centers or communication routes will facilitate the use
of a lake for commercial purposes. When selecting other real estates for comparison, the
aim is to find ones which will be characterized by similar general location. If this condi-
tion is met, general location will not require any further analysis – within the selected
group of lakes it will no longer be a factor which could differentiate transaction prices.
Detailed location in the appraisal of lakes mainly defines the communication acces-
sibility, such as the possible access to a lake on foot or by car. In extreme cases, a lake can
be inaccessible (by the power of an administrative decision periodically prohibiting entry
to state forests). Due to land configuration, access to a lake in a motor vehicle might be
impossible. If feasible, the quality of access roads should be specified. Surfaced, asphalt-
ed roads raise the attractiveness of a lake. They make fishery easier and offer more op-
portunities for organizing recreational services, e.g. angling. Main roads (administered
by the Province) which are cleared from snow first, will be valued most highly; next are
roads overseen by the local administrative district (powiat) and finally surfaced roads for
local traffic. Accordingly, lack of a surfaced road is valued very low.
The above states (degrees) of one attribute (communication access) can be assigned
a qualitative scale, e.g. access conditions by car very good, good, average, not good, bad,
a scoring scale from 5 to 1, or a relative scale, reflecting other than linear relations be-
92
tween the consecutive states, e.g. 5,3,2,1 and 0. The choice of a scale should be justified
by previous studies on the effect of this attribute on shaping transaction prices for lakes.
The quality and trophic state of a lake are among the basic attributes which most often
determine fishing, angling, recreational and ecological values of lakes. In compliance with
the current legal regulations, there is a five-grade scale for evaluation of the quality of
water (Ordinance of the Minister for the Environment of 20 August 2008 on classification
of the status of consolidated parts of surface waters, JoL No 162, item 1008). The most
desirable are class one waters, but these are very rare in Polish lakes, or class two waters,
because they ensure that the lakes can be fully used for recreation or angling. Although
the law does not state the suitability of class three waters for fish culture, such waters
are often inhabited by cyprinid fish, including many attractive game species. Lower water
quality classes, four and five, practically disqualify a lake as a potential site for recreation.
In the practice of lake valuation, the limnological system of lake classification is use-
ful, as it combines the trophic state (fertility) of lakes with the degree of their devel-
opment advance. In this system, oligotrophic, mesotrophic (α- and β-mesotrophic) eu-
trophic and dystrophic lakes are distinguished (Starmach 1976). The most desirable ones
are oligotrophic lakes with clean waters, which ensure full recreational use of a water
body. However, all lakes undergo eutrophication. Under natural conditions, this process
is irreversible, and the existing lake reclamation methods, which are to restore the origi-
nal character of lakes, are costly and seldom completely successful.
The character of a drainage basin and the way it is used can either pose certain threats
or create some protective zone around a lake. The main types of lakes’ immediate catch-
ment basins, in the order from the least to the most obtrusive ones, are: forests, meadows
and pastures, arable lands, recreational areas, rural and urban areas, industrial areas.
Forests protect lakes, counteract erosion and reduce amounts of biogenic substances
in surface runoffs. Forestry can create certain limitations and risk. It has already been
mentioned that entry to a forest can be prohibited periodically. Other unfavourable ef-
fects can consist in dumping timber processing waste to lakes, storing timber in water,
or felling and transporting cut tress near the shores of a lake.
Grasslands, meadows and pastures are characterized by relatively low farming intensity,
including a low level of fertilization and application of pesticides or herbicides. Grasslands
prevent water erosion and reduce amounts of biogenic substances in surface runoffs. With
regard to pastures, their negative effect on lakes depends on the intensity of animal rearing
and grazing. A large herd of farm animals near a lake could be a threat. It most often causes
complete damage to the shore and littoral zones of the lake where animals are watered.
Arable lands are an unwanted element in a lake’s immediate basin. To what extent
they can threaten a lake depends on the intensity of plant production, application of
mineral fertilizers, cultivation techniques, etc.
The recreational use of lands surrounding lakes certainly enhances their mutual at-
tractiveness. There is positive feedback – recreational use of land raises the commercial
value of a lake, and the presence of a lake determines the specific character of the sur-
rounding land. There is just one conditions – lakes must be unpolluted so that they can
be fully employed for recreational use.
93
Recreational facilities situated in a lake’s basin – tent and camping sites, summer
houses and guesthouses –turn into populated areas during the tourist season compa-
rable to urban settlements. They are often accompanied by some developed infrastruc-
ture, including roads, car parks, marinas, shops, bars, etc. If such areas are not sewered
and the permissible norms of loads related to the recreational capacity of lakes and their
ability to assimilate biogenic substances and pollutants are not respected, the lakes are
likely to be degraded.
Rural settlements with no sewage systems lying in basins of small lakes are the
main cause of their degradation. In a very short time, wastewater and sewage dis-
charged directly from households and farmsteads into a lake as well as liquid manure
sprayed over fields can turn a lake into a water reservoir usable just for irrigation in
farm practice. Sewered rural settlements and, more distinctly, urban ones can also
accelerate eutrophication and degradation of lakes. By developing land along lake
shorelines, reinforcing the banks, building concrete or tarmac paths along shores, con-
structing marinas, piers, etc., we devoid a lake of its natural protective zone, increase
the rate of surface runoffs and decrease the self-cleansing capacity of waters, which
is already impaired.
A non-degraded lake in a recreational sphere of a town – in a park or a wood – is an
extremely attractive component of this area, as it makes its open space larger and more
varied. Depending on its size and quality of water, such a lake can be used for organizing
a bathing area or a water marina, etc.
Lakes, even small ones, rarely have a basin that is used in a uniform manner. As
a rule, one form of use, e.g. forest or grassland, is accompanied by other uses, e.g. ar-
able land, rural buildings, and infrastructure facilities – railroad embankments, roads,
electrical power lines, etc.
When we have a sufficient amount of data available, we can evaluate the lake’s basin
and its effect on the lake using specialist methods. In the method suggested by Bajkie-
wicz-Grabowska (Bajkiewicz-Grabowska and Malczyk 1988), a lake’s basin is assessed
as a supplier of potential matter and a lake as its recipient. For evaluating the degree to
which the basin affects the lake, the following parameters are employed:
- lake coverage index (quotient of the total area of the basin and surface area of
the lake),
- the water balance type of the lake (lakes with surface inflow and outflow, no
outflow and with outflow),
- the degree to which the lake’s immediate catchment area lacks outflow,
- the average slope of the immediate basin,
- permeability of the ground,
- type of use of the lake’s immediate basin.
Each of these parameters is evaluated on a 0 to 3 point scale, where 0 means a very
weak effect on mobilization of matter in the basin and lack of supply of matter to the lake,
and 3 stands for very large potential of matter mobilization and quick transport to the
lake. The value of an arithmetic mean of the number of points scored from the assessment
of each parameter enables us to classify a given basin into one of the four groups:
94
- group I (x<1.0): the basin practically shows no ability to supply matter to the
lake because of the whole set of physicogeographical properties which strongly
reduces surface runoffs,
- group II (1.1 < x < 1.4): the basin shows little ability to mobilize the area load of
matter, which can hardly reach the lake,
- group III (1.5 < x < 1.9): the basin is moderately able to mobilize the matter and
supply it to the lake,
- group IV (x > 2.0): the basin has good ability to mobilize the area load of matter
and transport it to the lake.
The natural tolerance of lakes to effluents from their catchment areas is also evalu-
ated on a 0 to 3 scale, with 0 indicating high tolerance and 3 – no tolerance. The follow-
ing properties of a lake are evaluated:
- the average depth of a lake,
- the value of the quotient of the lake’s capacity and the length of its shoreline,
- % of water stratification,
- the value of the quotient of the lake’s bottom within the epilimnion and the ca-
pacity of the epilimnion,
- Schindler’s index (the quotient of the total area of the drainage basin and the
lake’s volume),
- The average annual water exchange intensity (the quotient of the average annual
outflow from the lake and the lake’s capacity).
The value of the arithmetic mean derived from the points scored on each of the lake’s
tolerance parameters will classify a given lake into one of the four categories of toler-
ance (Bajkiewicz-Grabowska and Malczyk 1988);
- category I (x<0.8): lake highly tolerant to effluents from the catchment area,
- category II (0.9 < x < 1.6): lake moderately tolerant,
- category III (1.7 < x < 2.4): lake not very tolerant,
- category IV (x > 2.4): lake strongly dependent on the catchment area and not
tolerant to its effluents.
The total valuation of the natural vulnerability of a lake to supply of matter and its
tolerance to effluents from the catchment area helps to distinguish four catchment-lake
systems, which, in lake valuation, reflect four states of the attribute type of the catch-
ment and threats:
System I is such a basin-lake system, in which both the natural properties of the lake
and those of its basin do not favour eutrophication of waters. The lake is tolerant to
external influence and its catchment area is not very active in supplying potential matter
to the lake.
System II is a system, in which the conditions in the catchment area unfavourable to
the lake (high possibility of mobilizing the potential area load of matter) are counterbal-
anced by the high natural resistance of the lake.
System III is composed of favourable conditions in the basin (the basin is not very ac-
tive in supplying potential matter to the lake) co-exist with the low tolerance of the lake.
95
In this case, any interference in the catchment conditions will quickly lead to excessive
eutropication of the lake.
System IV is a very unfavourable set of conditions, which lead to rapid eutrophication
of the lake. The natural properties of the basin favour runoffs and the lake lacks natural
tolerance of external influence.
Kudelska et al. (1991) distinguish three categories of lake vulnerability to anthropo-
genic effects, depending on lakes’ formo-morphometric, hydrographic and basin-related
characteristics, which – in combination with three lake purity classes – form six possible
states for total evaluation of the quality of water, trophic state, basin and threats to
a given lake, as well as possibilities of lake reclamation, depending on the water purity
class and category as well as vulnerability to man-made influence. The natural features
of the lake and its basin are decisive factors shaping the quality of waters in lakes under
a relatively low anthropogenic pressure. With regard to lakes which receive wastewater
and sewage, this harmful human activity is the basic cause of degradation of these wa-
ter bodies, regardless of their natural conditions.
The landscape values ascribed to a lake are a product of the mutual action of a set
of factors, among which the important role is played by the shape of a lake, the course
and development of its shoreline and the lake’s surroundings. The feedback relations be-
tween these factors mean that the evaluation of their aggregate effect usually diverges
from the sum of isolated, single evaluations of each of the parameters.
Smaller lakes, up to 10 ha in area, oval or round ones and lying in land depressions
or among hills, are not less attractive although their shoreline is poorly developed. But
among larger lakes, the ones which are irregular in shape and elongated are more at-
tractive in terms of landscape-related evaluation. The more developed the lake’s shore-
line, and the more irregular its shape, the more inlets, peninsulas or islets it possesses.
A measurable index of the shoreline development is the quotient of the length of
the shorelines of the lake and its islets (L) and two radicals from the product (π) and the
surface of the water table (P):
−1
/2/ Wl = L ⋅ (2 ⋅ p ⋅ P)
The index of the shoreline development tells us to what extent the shoreline of a giv-
en lake is divergent from the base model of a circle. A lake that is an ideal circle in shape
is characterized by the index (Wl) equal 1. As the value of this index increases, the shore-
line is gradually more and more developed (Filipiak et al. 1995):
• 1.0 to 1.5 – the shoreline poorly developed,
• 1.5 to 2.0 – the shoreline moderately developed,
• 2.0 to 3.0 – the shoreline well developed,
• over 3.0 – the shoreline very well developed.
The varied landscape around the lake enhances its attractiveness in terms of land-
scape values. An examples of attractive lake surroundings is a mixed forest with trees
of different age. Country cottages or summer houses skillfully designed to fit the forest
surrounding can also create interesting landscape compositions.
96
Elements of infrastructure like railroad and road embankments, visible high-voltage

lines, pipelines, etc., usually have a negative effect on landscape values. An exception to
this rule could be bridges and flyovers as well as water management or inland naviga-
tion facilities, e.g. locks, constructed in the old, historical times and frequently endowed
with the status of a monument of history. Another example of an attractive component
of the landscape surrounding lakes are former water towers or bunkers dating back to
the times of World War Two.
The recreational attractiveness of lakes depends on the forms of spending leisure
time which are possible in each case. The main forms of recreation are:
• weekend and holiday stays,
• water sports and sailing,
• angling.
Weekend and holiday stays are either individual or organized stays by a lake, for
a weekend or longer, in agritorusim farms, holiday centres, hotels, guesthouses or
camps, etc. This form of tourism mainly involves bathing, swimming or making walks
and trips in the nearby area.
Water sports and angling can also be pursued as part of holiday stays, but owing to their
specific character, these two forms of recreation have been presented as a separate entity.
Lakes suitable for bathing, swimming, water sports and other recreational aims must
belong to at least class two of the water purity classification (acc. to the Ordinance of
the Minister for the Environment of 20 August 2008 on classification of the status of
consolidated parts of surface waters, JoL No 162, item 1008). Bathing in lake water is
most often combined with sunbathing. Therefore, the most suitable lakes are the ones
which have sections of the shore gently sloping into the lake, with natural beaches hav-
ing direct access to water. The bottom of a lake should be hard, most preferably covered
with sand or sand and gravel. If the lake’s bottom is muddy and overgrown with emer-
gent plants, then jetties and piers can give access to water.
Another typical activity undertaken as part of holiday stays by a lake are walks and
hikes in the nearby area. Whether a lake is suitable for this form of recreation depends
on the landscape values of the lake itself and the surroundings. Trails of walks and hikes
can be prepared in many ways – natural or hardened surfaces, foot or bike paths.
Water sports include rowing, canoeing and other forms of water recreation with the
use of boating equipment, except sailing. Motorboating, very attractive although ecologi-
cally questionable, can also be considered, with such equipment as motor boats, water
skis or water scooters, etc. Unfortunately, motorboating is often a component of luxury
water recreation offered by holiday centres of the highest standard, in which civilization
and technical amenities are appreciated as much as natural values of the environment.
Relaxing trips in boats and other similar water equipment do not require very big lakes,
but on smaller lakes there is a risk that interests of different lake users could collide. Canoe-
ing does not need large water reservoirs, provided the ones available are connected with
one another, creating canoeing trails. This condition is quite difficult to fulfill in the case of
lakes classified as stagnant water bodies. By analogy, sailing is a form of recreation typical
of large lakes, which are flowing waters. Windsurfing is the least demanding water sport
97
in terms of the size of a lake. It can be presumed that a lake around 10 ha in area can be
used for windsurfing as long as it is elongated and has bare shores.
Angling is one of the main recreational uses of lakes, which frequently shapes their
commercial value. The angling usability of a lake mostly coincides with their fishing us-
ability (Szczerbowski 2008). This coincidence can be justified by the fact that landscape-
related and other values of lakes, important for anglers but of secondary interest to
fishers, are submitted to a separate evaluation of lake appraisals.
Table 3. Evaluation of the grades (states) of market attributes of lakes on market A.
Attribute Description Grade

Access by regional road, dirt road to the lake no loner than 1.0 km;
area attractive or potentially attractive for recreation (varied land-
scape, forest, infrastructure), possible walking or cycling tourism;
lake basin managed in accord with lake protection principles, with good
no intensive farming or industries, without residential settlements
with no sewers; local people hospitable and friendly.
Location, land- Access by local surfaced road, dirt road to lake no longer than
scape-related 1.0 to 5.0 km; area of average recreational and landscape assets;
values moderate
no special threats causing the lake’s more rapid eutrophication
found in the basin.
Difficult direct access to lake; periodical bans on travel (e.g. state
forests, private grounds), boggy land, damaged road; unattrac-
tive surroundings, uninteresting landscape; basin intensively
bad
used for farming, with industrial buildings, summer houses or
households with no sewers; areas of high unemployment, local
people distrustful or unfriendly
Clean waters, with no evident blooms in summer, bottom mostly
covered with sand and gravel’; very good swimming and sun- good
bathing conditions
Quality of wa-
Moderately clean waters, with regularly occurring blooms of
ter, including
phytoplankton, moderately intensive in summer, bottom mostly
possible crea- moderate
hard, sandy and muddy; sufficient conditions for swimming and
tion of bathing
sunbathing
beaches
Fertile waters, low transparency, with intensive algal blooms in
summer, water quality inadequate to set up bathing beaches bad
(below class II), muddy, sinking bottom
Good – abundant fish valued by fishers and anglers, including
good
the fish obtained from fish fry releases
Lake average in terms of fishing and angling value, under heavy
Fishing and moderate
poaching pressure
angling assets
Lakes with big oxygen deficits, causing mass death of fish, close
to dystrophic lakes, heavily polluted or threatened with waste- bad
water and sewage, depressing food value of fish
Source: Turkowski 2003
In the evaluation of the fishing usability of lakes, the lake fishing index (Wr) (Filipiak
et al. 1995) can be helpful. The index is the ratio of the lake’s shoreline length (L) to the
water table surface area (P):
/3/ Wr = L ⋅ P −1
98
The rising value of the fishing index usually means a more developed littoral zone of
the lake, which in turn indicates a higher number of spawning and foraging grounds.
When forecasting the value of lakes whose dominant function is angling, a synthetic
measure of lake suitability for angling can be engaged, which is composed of the natu-
ral conditions (the ratio of the shoreline development, emergent vegetation, access to
the water table and to the water body) and the lake’s angling management (construc-
tions raised in the lake shore zone, the volume and quality of fish releases) of the lake
(Skrzypczak 2005).
For an appraisal of a lake as a real estate, first of all, we take into consideration these
lake properties which can directly transform into actual financial profits to potential lake
owners. In this context, the most important are the fishing and angling usability as well
as the recreational values of lakes. Location, trophic state and potential threats to a lake
largely determine the attainment of the benefits mentioned earlier. The selection and
criteria for the valuation of attributes should be adjusted and modified depending on
the conditions on a given market (Tabs 3 and 4).
Table 4. Evaluation of the grades (states) of market attributes of lakes on market B.
Attribute Description Grade

close proximity (up to 5 km) to large or medium-sized ag-
glomerations or important routes, convenient access by
tarmac road or no more than 0.5 km by dirt road; the land
adjacent to the lake belongs to the whole real estate; very good
distance to above urban agglomerations or important

Location, access routes above 5 km to 10 km, access by dirt road above
0.5 to 1.0 km; the land adjacent to the lake belongs to the
moderate
whole real estate;
access as above but restrictions due to roads located within

state forests; distance to an urban agglomeration or main
routes over 10 km, access by dirt road above 1.0 km. not very good
attractive recreational or potentially recreational grounds
(varied landscape, forest), no intensive farming or indus-
try, either no or unobtrusive land development; good
Nature and land- intermediate between good and not very good character-
scape values istics; moderate
surroundings dominated by an agricultural catchment

area or developed land; pollution or potential pollution
from farming, households or industries; not very good
lakes with no oxygen depletion causing mass death of fish,
fertile or very clean, deep; very good
fertile or not very eutrophic lakes, deep and not over-

Fishing values good
grown with plants;
shallow lakes, overgrown with plants, and/or with oxygen

depletion causing mass death of fish not very good
Source: Turkowski 2008
99
Analysis of the market of similar real estates

Such analysis begins by reviewing lake buy/sell transactions on the local market over
the past two years. When too few transactions are found, the area and time period
searched are extended. Consequently, we should be able to identify transactions ap-
propriate for the location of the lake being appraised, conditions under which they were
concluded, selling prices and dates of concluding transactions and the attributes of sold
lakes. Because of different dates on which lakes were sold, it is necessary to identify how
prices were shaped in time. If prices tended to rise or fall in time, they need to be cor-
rected accordingly to the same time period – the date when the value of the appraised
lake is determined (Tab. 5).
Table 5. An example of a comparison of lake buy/sold transactions.
Fish-
Date of Total real Corrected Location, Nature and
No of Price ing and
transac- estate area price access landscape
transaction PLN/ha angling
tion in ha PLN/ha conditions values
values
1 06.1997 4.02 3606 24949 not good moderate very good

2 06.1997 5.91 4653 29909 good good good
3 01.1997 4.67 3426 29749 good good very good
4 12.1998 7.47 2800 25032 not good moderate very good
5 12.1999 8.30 3723 23821 not good good very good
6 10.2001 3.78 13228 29235 good moderate good
7 04.2006 3.73 21716 28119 moderate not good moderate
8 04.2009 6.59 29393 29393 good moderate very good
Source: Turkowski 2009 modified
The next step in the valuation is the specification of the weight of attributes, that is
their percentage share, on shaping transaction prices. For this purpose, the lakes which
were bought and sold on the market are set in pairs so that they should be maximally
different in respect of just this feature for which the weight is calculated (Tab. 6). The cal-
culation is based on the assumption that the difference in transaction prices for the pairs
of lakes thus set was caused by the difference in the status of the analyzed feature:
C w − Cm
/4/ wi =
∆C
where:
wi – (i) feature index; having calculated indices for all attributes, it is converted to
a percentage according to the formula: ,
Wi
Wi (% ) = ⋅100%
∑ wi
100
Cw – unit price of a real estate of the highest status of the attribute (the best for this
feature),
Cm – unit price for a real estate of the lowest (worst) status,
Cmax. – maximum unit price,
Cmin – minimum unit price,
∆C – difference between the maximum and minimum corrected price in the collected
sample of market transactions on lakes.
Table 6. Calculation of weights of attributes of lakes submitted to market turnover.
No Attribute Prices for real estates in pairs wi index

C3 = 29749 0.97
C5 = 23821
1 Location, access conditions
C8 = 29393
C4 = 25032 0.72
On average 0,84
C2 = 29909
C6 = 29235 0.11
2 Nature and landscape values
C3 =29749
C8 = 29393 0.06
On average 0,08
C4 = 25032
C1 = 24949 0.01
3 Fishing and angling values
C8 = 29393
C6 = 29235 0.03
On average 0.02
Establishing per cent weights:
0.97+0.08+0.02 = 1.07;
Location, access conditions: 91%
Landscape values:7%
Fishing and angling values:2%
In order to keep to the minimum accidental results, calculations are completed for
at least two pairs (Tab. 6). In the given example, fishing and angling assets are assigned
a surprisingly low weight – just 2% effect on the level of buy/sold prices (Tab. 6). This
result should not be treated in absolute categories – it does not mean that these two
values are of low importance. It only means that the fishing and angling values were not
a feature which differentiated considerably the prices of lakes on the analyzed market.
If most of the real estates are similar in a given feature (e.g. fishing and angling values
are good or very good), this feature ceases to be the main reason why prices for real
estates are different. Once the per cent weight is multiplied by ∆C (formula 3), the value
of transaction price corrections is obtained, expressed in an amount in PLN/ha. When fix-
ing a price correction, we take into account the difference in a given feature (attribute)
expressed in grades. If these differences are at extreme ends, e.g. location of and acces-
sibility to one of the compared lakes are good but poor with regard to the other one,
101
the correction will be 100%, but if they are poor and moderate, then it will be 50% on
a three-grade scale (Tab. 7).
Table 7. Per cent and amount ranges of corrections assigned to particular market attributes of lakes.
Amount of correction
Per cent of weight
Attribute Evaluation in PLN/ha
(%)
(*weight)
not good 0 0
Location, access moderate 50 2922
good 100 5345
not good 0 0
Natural and land-
moderate 50 213
scape values
good 100 426
not good 0 0
Fishing and angling
moderate 50 61
values
good 100 122
Having computed the corrections among the collected lake purchase transactions, at
least three are chosen, for which the lakes were the most closely similar to the one being
appraised. The objective is to find an answer to the question: what would the price for
each of these lakes be, if the grades (states) of all their features were the same as the
ones assigned to the analyzed lake? The answer demands that the transaction prices be
corrected by the corrections presented above. For example, if the nature- and landscape-
related values of a lake, whose transaction price is known, were good on the selling
date and those of the appraised lake just average, this would mean that its hypothetical
price would be less by half the weight of that feature, i.e. by 213 PLN/ha (Tab. 8). Other
features are compared analogously.
Table 8. Per cent and amount ranges of corrections assigned to particular market attributes of lakes.
Comparative real Amount range

Appraised real Weight of Correction
Attribute estate Lake no 2 (ΔC * weight)
estate Lake x attribute (%) (PLN/ha)
(tab.5) in PLN/ha
0 0
Location,
good good 50 2922 0
access
100 5345
Nature and 0 0
landscape moderate good 50 213 -213
values 100 426
Fishing and 0 0
angling val- good very good 50 61 - 61
ues 100 122
Sum - 274
The corrected price obtained from selling lake no 2, i.e. 29,909 PLN/ha (Tab. 5) is
enlarged by the sum of the corrections equal 274 PLN/ha (Tab. 8) and so a hypothetical
price of the appraised lake is attained, such as 29,635 PLN/ha. By analogy, we correct
transaction prices from the other two lake buy/sell transactions (6 and 8, Tab. 5). The
102
market value of the appraised lake which we have searched for is eventually an average
of the three transaction prices corrected as explained above:
x2 x6 x8 29635 29235 29332

W , W 29401 PLN/ha
3 3
Having accounted for the size of the lake, this value is:
29 401 PLN/ha * 17.8 ha = 523 338 PLN.
The level of transaction prices of undeveloped farmland real estates on the local
market, proper for the valued lake, was on average half that price, in terms of the same
area unit. A higher value of a lake versus arable land can be attributed to the lake’s
recreational value as well as to the fact that lakes are rarely offered for sale on the real
estate market.
Lakes are very special and unique ecological systems, in which plant and animal
populations dwell on different trophic levels; there are many endemic species or the
ones which are becoming extinct. Their importance in commercial valuations of lakes is
secondary, that is they matter only if they have some influence on the use features of
lakes. However, if owing to some particular ecological values of a lake, e.g. occurrence of
rare plant or animal species, the lake is given some status of a conservation area, then in
market relations this is usually a factor which lowers the value of the lake. In some cases,
when the owner of a lake covered by a nature conservation plan is eligible to compensa-
tion for lost revenue, then the market value of this lake, which reflects the market value
of the ecological assets, will be proportional to the amount of compensation and the
period of time for which it is paid.
Summary
Regardless of their use values, the value of lakes as real estates largely depends on
what approach, method and techniques of valuation will be chosen. Thus, the choice
should be justifiable in accordance with the purpose of the appraisal, type, location and
intended use of a real estate in spatial development, technical facilities it is equipped
with, frequency of concluded transactions involving similar real estates, possibilities of
acquiring information on the range of transaction prices as well as the conditions under
which those transactions were done. The main market analyzed for the purposes of real
estate appraisals is the local market, which covers transactions from the past two years.
However, due to a very specific nature of the turnover of lakes, this area and time period
are most often extended.
Valuation of lakes as real estates is connected with the valuation of rights that poten-
tial owners may acquire. As a rule, the value thus estimated does not reflect the value of
a lake as a natural resource. Such values go beyond the issue of ownership rights and - in
extreme cases – beyond the scope of human uses. These questions, however, belong to
the field of natural resources appraisal and require a separate presentation.
103
References
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zagrożenia jezior eutrofizacją na przykładzie kilku jezior Pojezierza Gostyńskiego. In: Churski Z.
(red.) Naturalne i antropogeniczne przemiany jezior i mokradeł w Polsce. Materiały Konferencji
Komisji Hydrograficznej PTG w Bachotku, Toruń.
Falkowski S. 2006. Gospodarka rybacko-wędkarska w zbiornikach wodociągowych na
przykładzie zbiornika zaporowego Goczałkowice. In: A. Wołos. (Ed.). Rybactwo, wędkarstwo, eko-
rozwój, Wyd. IRŚ, Olsztyn: 135–149.
Filipiak, R. Trzebiatowski, J. Sadowski. 1995. Rybactwo-przewodnik do ćwiczeń z przedmiotu
Gospodarka na wodach otwartych. Wyd. Akademia Rolnicza w Szczecinie.
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Kudelska D., Cydzik D., Soszka H. 1991. Monitoring podstawowy jezior Polskich. Gospodarka
Wodna, 6:127-130.
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nomics and Management 32: 22–37.
Skrzypczak A. 2005. Ocena przydatności rekreacyjnej naturalnych zbiorników wodnych dla
wędkarstwa. Folia Turistica, 16: 115–129.
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PWN, Warszawa: 336-424.
Szczerbowski J. 2008. Rybactwo śródlądowe. Wyd. IRŚ Olsztyn.
Turkowski K. 1994. Problem ekonomicznej oceny wartości zasobów wodnych. In: Łojewski S,.
Czajkowska J. (Eds.). Materiały pokonferencyjne, tom I Ogólnopolskiej Konferencji „Ekonomika za-
sobów wodnych i ochrony wód”. Bydgoszcz – Warszawa: 33–41.
Turkowski K. 2003. Wycena wód i gruntów pod wodami. Wyd. Educaterra, Olsztyn.
Turkowski K. 2008. Wycena jeziora Krzywek. Ekspertyza wykonana w Katedrze Rybactwa Jezio-
rowego i Rzecznego, UWM w Olsztynie, na zlecenie SITR sp. z o.o. w Olsztynie, maszynopis.
Turkowski K. 2009. Wycena jeziora Pogorzelec. Ekspertyza wykonana w Katedrze Rybactwa Je-
ziorowego i Rzecznego, UWM w Olsztynie, na zlecenie Sądu Rejonowego w Sejnach, maszynopis.
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Cieśla & A. Lirski (eds.) Multifunctionality in Pond Aquaculture in Poland. Current Status. Warsaw
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Legal regulations
Journal of Law of 2010. No 102, item 651. Act of 21 August 1997 on real estate management
(consolidated text, , with amendments). in Polish
Journal of Law of 2005. No 239, item 2019 Act of 18 July 2001- Water Law (consolidated text,
with amendments). in Polish
Journal of Law No 207, item 2109 Ordinance of the Council of Ministers of 21 September 2004
on real estate appraisal and drawing valuation surveys (, with amendments) in Polish
Joural of Law No 162, item 1008 Ordinance of the Minister for the Environment of 20 August
2008 on on classification of the status of consolidated parts of surface waters,).
104
Fishery types of lakes in the
context of lake eutrophication
and changes in the structure of
ichthyofauna
Andrzej Skrzypczak1, Andrzej Mamcarz1,
Adam Gierej2
Department of Lake and River Fisheries,

University of Warmia and Mazury in Olsztyn1
Polish Angling Association, Region in Ciechanów2
e-mail: andrzej.skrzypczak@uwm.edu.pl
Abstract
The relevant references have been reviewed in order to present the taxonomy of lakes
according to the ichthyofauna which inhabits these lakes. Fishery types of natural water
bodies have been discussed alongside their morphological features and morphometric
indices of lakes within the fishery types. Changes over time occurring in the structure of
the exploited ichthyofauna in Polish lakes in the context of the adopted fishery lake type
classification have been analyzed, paying particular attention to the events taking place
in the lakes in north-eastern Poland. The question of the lake classification according to
the fishery type has been discussed against the background of the changes which take
place in fish assemblages due to lake eutrophication and succession mechanisms. It has
been indicated that the functionality of the fishery classification of lakes is limited due to
its static nature. This limitation can be overcome by including the concept of a dynamic
transfer of undisturbed and stratified water bodies classified as vendace type and bream
(α) type into bream (β) type of lakes, being in a different state of disturbance depend-
ing on the trophic state and the specific oxygen conditions. The authors emphasize the
need to conduct further observations of potential transformations of fish assemblages
in Polish lakes because of the changes in the model of fishery management of natural
water bodies and the increasing role of lakes in tourism and recreation.
Introduction
Lakes are thought to be short-lived forms in the context of the Earth’s geological life.
The basic system of classification of natural aquatic bodies relies on their genesis. Most
105
of the lakes in the northern hemisphere have post-glacial origin and appeared several
thousands of years ago when mountain glaciers and ice sheets shaped the surface of the
earth. The variety of lakes’ morphological shapes, mainly the shape and depth of lakes’
bowls, is a direct reflection of the specific nature of glaciations processes. The most
popular types of lakes are ribbon, moraine and thaw lakes (Choiński 2000).
Lakes are subjected to the mechanism of succession, whose rate and results are
a product of a number of internal and external factors as well as some complex process-
es which take place in their ecosystems. Consequently, even when their morphological
properties are similar, lakes are inhabited by a variety of plant and animal communities;
they also show some specific characteristics of the chemical and biochemical processes
engaged in matter cycling and energy flow. The above features have been used to de-
velop systems of lake classification. The proposed taxonomies based on limnological and
ecological properties of lakes differ from one another in their complexity and specific na-
ture of suggested indices. Apart from thermal classifications or mictic types, another cri-
terion for lake divisions could be, for example, the composition of crustacean plankton
(Bowkiewicz 1935, Lityński 1935) or characteristic communities of fauna (Bernatowicz’s
typology, 1960). Among the most popular typologies is the one developed by Neumann
and Thienemann, which was proposed in the early 1930s, and the key to dividing lakes
into categories was their trophic state. The main advantage of this classification is that
it is compatible with the natural succession of water bodies; another one is the continu-
ity of transfer through subsequent stages within the distinguished types and subtypes.
An example of the application of Neumann-Thienemann’s typology to post-glacial lakes
situated in central Europe is the division made by Stangenberg (1934), modified by
Wiszniewski in 1953 (Choiński 2000). This is why today we use, rather intuitively, the
division of lakes into oligo-, meso- and eutrophic ones, with all ‘hues’ of eutrophic states
between. At the same time, we can hypothesize that the limited usability of lake clas-
sifications based on presence of absence of live organisms is due to the complexity of
their applications, but in many cases also because some of the classification systems
are very static. A question then arises – can the composition of ichthyofauna in natural
water bodies serve as a basis for lake typology, whose functionality will not be limited by
changes taking place in lakes’ ecosystems?
Fishery and ichthyologic classifications of lakes

The oldest system based on leading fish species is the classification suggested in 1881
by Max von dem Borne and developed for Alpine lakes (Korycki 1976). However, the major
contribution into the development of fishery types of lakes was made by Somov, who
carried out observations on lakes situated in the European part of Russia and in 1920 sug-
gested a division of lakes which characterized their ichthyofauna and environmental con-
ditions (Somov 1920). In the 1930s, as the limnological studies on the typology of Polish
lakes progressed, an attempt was made to classify them according to the structure of the
fish communities in given lakes (Lenkiewicz 1936, Rudnicki 1947, Sakowicz 1950). The
question of lake classification was then considered to be a key problem in limnology and
in the then developed system for the fisheries management of Polish natural water bodies,
especially after World War Two (Bernatowicz 1949, Backiel and Zawisza 1956). Based on
106
Fishery types of lakes in the context of lake eutrophication and changes…
the experiments run by Somov and transferring their results to the specific conditions of
the Polish lakes, Staff suggested his own typology of lakes (1950), In the mid-1950s, it was
assumed that studies on a general outline of the limnological lake typology had finished,
although a series of difficulties related to the synthetic nature of this problem were indi-
cated (Wiszniewski 1953). Nonetheless, an important step in the work on developing fish-
ery classification of lakes was the typology suggested by Abrosov (1957), who improved
the division suggested earlier by his predecessor, another Russian scientist, Somov. Con-
sidering the work on developing the natural foundations for the lake management system
in Poland, the attempt made by Stangenberg (1956), who tried to combine the results on
the limnological classification of lakes with their fishery characterization is an extremely
valuable contribution. The fishery typology of Polish lakes, developed and implemented at
the turn of the 1950s and 1960s by the Central Fisheries Management (Centralny Zarząd
Rybactwa, the CZR), was based on the parameter of a lake’s depth and some selected hyd-
robiological characteristics (Rudnicki et al. 1971). Compared to the previous classifications,
this one was much simpler (Tab. 1).
Table 1. Fishery classifications of lakes.
Fishery typology according to

the CZR
Somov (1920) Staff (1950) Abrosov (1957)
(Rudnicki et al. 1971)
lake trout trout lake trout vendace
common whitefish common whitefish common whitefish bream
bream vendace bream zander
- bream-smelt - bream-smelt - bream-whitefish tench-pike
- bream-bleak - bleak-bream - bream-vendace Crucian carp
zander perch-roach - bream-smelt
perch-roach Crucian carp - bream-bleak
Crucian carp tench
perch
perch-roach
pike
Crucian carp
The names of the five types distinguished by the CZR correspond to the fish species
which, owing to their biology and environmental requirements, find the most suitable
habitat conditions in given lake category. This division, binding until the present day, is
characterized by a reduction of subtypes within the category of bream lakes and an ad-
dition of the zander type of lakes, which among the earlier classifications was included
only by Samonov. An inclusion of zander, which appears numerously in lakes, in lake
typologies was thought to be troublesome. Staff abandoned the zander type of lakes
and treated the species as a typical riverine one. This problem is quite well-seen in the
classification of zander lakes elaborated by Bauch (1955), which divided them into four
main types, one of which was further broken into two subtypes (Korycki 1976).
Classification of lakes based on the leading species of fish is commonly referred to as
fishery typology. Such classification systems, by indirectly including several morphologi-
107
cal properties of natural water bodies and specific environmental conditions in which
aquatic reservoirs lie, encompass an ecological aspect as well. The ichthyologic character
of the suggested classifications is also supported by the fact that the key fish species
in the distinguished types do not have to appear most numerously but they are just an
indicator of the specific nature of a given lake and a base for drawing conclusions on
the species composition of the ichthyofauna community and the existing environmental
conditions. On the other hand, the above typologies also implied productivity of each
type of lakes by referring to the potential fishery capacity. In post-war Poland, the need
to develop a separate typology was driven by the fact that it was necessary to establish
uniform guidelines and directions for fishery management of lakes, for example in order
to create a proper fish brood release policy (Bernatowicz 1949).
Fishery typology of lakes versus their morphological characteristics

In the context of the changeability of physicochemical parameters and abundance of
lake waters in nutrients, and including the differentiated rate of these parameters becom-
ing more permanent, morphological properties and morphometric indices can be con-
sidered as relatively stable. Owing to their importance, they are a starting point for any
evaluation of the character of a water body in terms of water mixing, and indirectly its
thermal and oxygen conditions. This justifies the need to analyze differences in the lakes’
surface area, maximum depth, average depth, relative depth, depth index and shoreline
development ratio. Some broad analyses have been made in this respect for 2,078 lakes in
north-eastern Poland, which in total cover over 137,000 ha (Skrzypczak 2008).
Based on the analysis of morphometric parameters of 488 tench-pike type lakes, 318
bream type lakes, 139 vendace type lakes and 86 pikeperch type lakes, it was concluded
that the most specific characteristics can be found among vendace lakes, which differ
statistically from the other types in all the analyzed parameters (Tab. 2), The character-
istics of crucian carp type lakes, analyzed in the same category with small and shallow,
lakes, unclassified due to the limited availability of the data that would enable precise
classification as crucian carp type or tench-pike type of lakes, should be more carefully
verified. Several coinciding data and indirect properties indicate pikeperch type lakes.
The adopted division into fishery types of lakes is fully reflected by the parameters of
the maximum and mean depth. According to these parameters, the pikeperch type is
between bream type lakes and tench-pike type ones (Skrzypczak 2008). Mikulski (1964)
and Nagięć (1977) claim that all Polish pikeperch type lakes belong to the water bodies
which lack thermal stratification in the summer. It can therefore be presumed that, in the
light of the study reported by Patalas (1960), all bream and vendace type lakes are strati-
fied ones, described as III, IV or V degree of the static equilibrium. The pikeperch type
lakes in north-eastern Poland, in respect to the depth index and relative depth parameter
are closer to the tench-pike type. With regard to the shoreline development index, they
are similar to the bream type, but in respect of the surface area they do not differ either
from the vendace type of lakes.
Statistical analysis of the morphometric features of the lakes showed some devia-
tion of the variables of these features from the normal distribution, which necessitated
application of non-parametric tests (Łomnicki 2003). Analysis of skewness, by assuming
108
values different from zero, indicated the right asymmetry of the variables. This is also
confirmed by the observation of divergences between the arithmetic mean of the indices
and the value of the median, which is one of the measures of the central tendency, for
which half the observations (50%) lie above and the other half – below its value. The
results obtained suggest that derivation of the arithmetic mean of surface areas of lakes
and their maximum depths is burdened with the biggest error (Skrzypczak 2008).
Table 2. Diversification of morphological features of lakes in fishery types (after Skrzypczak 2008, changed).
Fishery type of lakes
and non-classified
crucian carp type
tench-pike type
pikeperch type
vendace type
bream type
Parameter
4.8 38.7 172.5 111.8 379.9

Mean ±SD (ha)
Area ±5.9 ±92.6 ±352.5 ±209.9 ±521.5
Median (ha) 2.9 A 18.7 B 95.5 CD 52.9 C 211.0 D
3.0 6.1 9.4 16.3 35.9
Mean ±SD (m)
Maximal depth ±1.9 ±3.9 ±6.4 ±6.9 ±13.1
Median (m) 2.5 A 5.1 B 7.9 C 15.4 D 34.5 E
1.4 2.7 3.7 5.8 10.8
Mean ±SD (m)
Mean depth ±0.8 ±1.5 ±1.5 ±2.1 ±3.9
Median (m) 1.2 A 2.5 B 3.6 C 5.7 D 10.2 E
0.487 0.427 0.448 0.374 0.311
Mean ±SD
Depth index ±0.128 ±0.094 ±0.118 ±0.106 ±0.078
Median 0.500 A 0.440 AB 0.450 AB 0.370 C 0.300 D
0.007 0.013 0.011 0.019 0.025
Mean ±SD
Relative depth ±0.004 ±0.009 ±0.011 ±0.011 ±0.015
Median 0.007 A 0.011 AB 0.008 AB 0.016 C 0.021 D
1.34 1.57 1.92 1.91 2.28
Shore develop- Mean ±SD
±0.17 ±0.41 ±0.67 ±0.67 ±0.69
ment index
Median 1.34 A 1.46 A 1.78 B 1.75 B 2.15 C
The values marked with the same letter index do not differ statistically (the Kruskal-Wallis test, P≤ 0.05).
Qualitative changes in the structure of exploited ichthyofauna

Observations of the results of long-term commercial fish catches often serve as a ba-
sis for drawing conclusions about the changes which occur within the exploited fish
communities. Analysis of fishery exploitation and its effects has been frequently con-
ducted in order to gain more in-depth knowledge of the species composition and ratios
between species in exploited fish populations (Leopold 1968, Skrzypczak 2008) or to
evaluate the changes that take place in whole lake ecosystems (Colby et al. 1972, Hart-
mann 1977, Leopold et al. 1986).
109
After World War Two, Poland developed a system of the fishery industry organization
for inland waters (Sakowicz 1947). The system comprised a unique way of keeping ar-
chives of the effects of fishery exploitation. In 1951, ledgers were introduced in the State
Fish Farms, in which annual catches of fish in kilos were recorded, divided into species,
commercial assortments, and months of catches; different fishery treatments were also
noted, such as fish stock releases. These ledgers now enable us to discuss the question of
the succession of fished species and possible reasons why some have disappeared while
others have come to dwell in our lakes (Nagięć 1978, Leopold et al. 1986).
Changes to the structure of the exploited elements of ichthyofauna result from
a number of processes and a crucial role was played by direct modelling of the fish for-
mations by fish stock releases and catches. For Polish lakes, the first step toward those
changes was to define the groups of species that are valuable from the economical point
of view and Stangenberg (1956) included the following fish into that group: bream, tench,
roach, pikeperch, whitefish, vendace, pike, and eel. Broadening the list with perch, smelt
and bleak formed a typical set of lake fish that appeared in various spices combinations re-
sulting from auto-regulatory and adaptive processes in biocoenosis. At the same time, the
author mentioned that the whitefish is sparse and of economical significance in several
places only, and the local position of the smelt and bleak was decided only by the possibil-
ity of their mass catches. Stangenberg (1956) claimed that despite small qualitative chang-
es, that composition had shown large resistance capabilities in the recent centuries.
When formulating general assumptions for the fishery management, the main spe-
cies requiring special economic treatment due to some difficulties in maintaining their
population in exploited lakes included: the tench, whitefish, vendace, pikeperch, pike, and
eel (Stangenberg 1956). Special treatment meant particularly fish stock releases of those
species and, to a lesser degree, supporting their natural breeding or taking into special
form of protection. What regards the exploitation procedures, some guidelines to support
or limit selected species of fish in lakes were defined. According to Leopold (1968), they
aimed at increasing the general effectiveness of catches in particular fishery types of lakes,
especially with respect to the valued species. Also, general directions for a possible restruc-
ture of ichthyofauna in particular types of lakes were defined (Rudnicki et al. 1971).
The fish stocks were operated on a basis of such formulated rules in many lakes in
1951 - 1994, including 1,049 lakes in north-eastern Poland. The observations of changes
that occurred in that time in the structure of the exploited ichthyofauna were conducted
by Skrzypczak (2008). The dynamics of those processes becomes visible when compar-
ing the share of 13 components in the structure (17 species altogether for crucian carp
and prussian carp, whitefish and peled, as well as introduced herbivorous species were
presented together) including the extreme periods of the analysis (Tab. 3). In 1951 -
1964, which is in the time of developing and implementing the fishery typology by the
CZR., the most visible differentiation of share in all the lake types was manifested by
the bream, perch, roach, and eel. In 1981 - 1994, such a differentiation was noticed for
Coregoninae, the bream and the eel. In those two periods, the status of differences and
similarities between all the categories remained unchanged for the crucian carp and
prussian carp, silver bream, bream, tench, pikeperch and eel. When comparing the share
of particular species in the structure of catches between the initial and the final period of
110
the analysis, an increase of that parameter was noticed for: Coregoninae, the bream, the
roach, the eel and the pikeperch (apart from the pikeperch type of lakes) as well as for
the silver bream in the tench-pike type and bream type of lakes. On the other hand, the
decrease of such a share affected the crucian carp and prussian carp, tench, perch, pike,
and pikeperch in the pikeperch type of lakes as well as the silver bream in the pikeperch
type and vendace type of lakes.
Table 3. Structure of the exploited ichthyofauna (mean percentage and ±SD) of the lakes in north-eastern
Poland in 1951 - 1964 (I) and 1981 - 1994 (II) together with the diversification of the species structure
in fish catches for the fishery types (after Skrzypczak 2008, changed).
Fishery type of lakes
tench-pike type
pikeperch type
vendace type
bream type
Components of
Years
ichthyofauna
I 0.03 ±0.02A 0.30 ±0.39A 1.87 ±1.64 13.63 ±4.97

Coregonus albula (L.)
II 0.03 ±0.03 0.35 ±0.22 3.09 ±1.43 17.95 ±2.91
Coregonus lavaretus (L.) I 0.01 ±0.01A 0.01 ±0.01A 0.05 ±0.08A 0.42 ±0.32
+ C. peled (Gmel.) II 0.30 ±0.30 0.18 ±0.08 1.34 ±0.54 3.10 ±0.92
I 1.94 ±0.81 5.11 ±1.62 3.87 ±1.31 2.83 ±1.13
Anguilla anguilla (L.)
II 4.83 ±1.31 9.73 ±1.83 9.40 ±2.48 8.34 ±2.14
I 0.53 ±0.28A 10.02 ±3.67 0.96 ±0.56 0.13 ±0.08A
Sander lucioperca (L.)
II 1.43 ±0.49A 7.32 ±1.29 1.68 ±0.71 1.24 ±0.71A
I 18.12 ±2.44 9.31 ±2.33A 9.56 ±1.83 9.41 ±2.10A
Esox lucius L.
II 8.85 ±1.77 3.58 ±0.79 5.02 ±0.61A 4.36 ±0.81A
I 6.73 ±1.87 7.76 ±3.67 9.67 ±2.51 9.63 ±3.51
Perca fluviatilis L.
II 3.62 ±1.28A 2.06 ±0.50A 4.86 ±0.59 4.71 ±0.70
I 17.18 ±4.13 4.13 ±1.54A 4.26 ±1.29 3.35 ±1.16A
Tinca tinca (L.)
II 6.57 ±2.23 0.83 ±0.34A 1.13 ±0.41 0.73 ±0.25A
I 5.48 ±1.69 0.85 ±0.88 0.65 ±0.45A 0.29 ±0.20A
Carassius sp.
II 3.42 ±2.58 0.57 ±0.37 0.25 ±0.25A 0.09 ±0.12A
I 19.13 ±5.83 32.09 ±5.29 20.27 ±2.56 14.16 ±1.78
Abramis brama (L.)
II 32.30 ±2.40 42.81 ±3.30 22.55 ±2.83 18.14 ±2.38
I 18.49 ±2.54 10.18 ±2.51 30.69 ±7.86 26.89 ±3.15
Rutilus rutilus (L.)
II 22.73 ±3.87A 19.57 ±2.07A 38.97 ±2.77B 34.53 ±2.78B
I 7.72 ±2.58 12.87 ±4.08 4.88 ±1.26A 3.34 ±1.13A
Abramis bjoerkna (L.)
II 8.24 ±2.27 7.95 ±2.79 5.62 ±1.33A 2.88 ±0.64A
I 0.79 ±0.66A 0.60 ±0.52 0.23 ±0.13A 0.07 ±0.09
Cyprinus carpio L.
II 2.81 ±1.99A 0.53 ±0.29 0.14 ±0.08A 0.04 ±0.02A
Ctenopharyngodon idella (Val.) I - - - -
+Hypophthalmichthys sp. II 1.26 ±0.85A 0.90 ±0.81A 0.92 ±0.82A 0.11 ±0.24
The records marked with the same letter index do not show a statistical differentiation of the share in the catches structure of
the listed elements of ichthyofauna between the analysed fishery types of lakes (the Kruskal-Wallis test, P ≤ 0.05).
In the majority of studies on restructuring the ichthyofauna in Polish lakes, based on

the analysis of the amount of the total fish catch (kg), the efficiency of fishery (kg/ha) or
111
the relative fish catch (kg/ha per month) (Skrzypczak and Mamcarz 2003), eutrophication
is considered to be the main cause of those changes, which is propelled by human activ-
ity in lake drainage areas and the development of the recreation functions of the lakes
(Bnińska 1984). According to numerous authors, a fast increase of the lake trophism
level and disturbances in the littoral affected in the most spectacular way the decrease
of fish catches connected with that zone, which is that of the crucian carp, tench and
pike (Bnińska 1985, Pyka and Zakęś 1991). As the dependencies between the influences
of tench stock releases on the amount of its catches was proved (Bnińska 1985), a mul-
tidimensional decrease of those procedures in the lakes of the north-eastern Poland was
shown, which comprised of the decreases in the frequency of the fish stock releases and
the amount of the doses used (Skrzypczak i Mamcarz 2006a). In the light of the carried-
out analyses, it was also shown that the speed of the tench decline is faster in the lakes
of the largest percentage of their littoral surface, which is in the conditions that occur
in the tench-pike type of lakes (Mamcarz and Skrzypczak 2006). Similar trends in the
amount decreases of stock releases were noticed for crucian carp (Skrzypczak i Mamcarz
2005b). However, it should be stressed that the population of that fish in the multi-spe-
cies sets is never large, especially when making intensive stock releases with predators or
in the conditions of a strong food competition (Zawisza and Antosiak 1961).
The changes in the status of the Percidae fish family in the lakes should be consid-
ered in the light of the decrease in the perch population that was not subjected to stock
releases (Leopold et al. 1986), and the increase in the number of pikeperch habitats
(Skrzypczak and Mamcarz 2005c). In the lakes of the north-eastern Poland, that phe-
nomena were of the most dynamic character in 1951–1965 (Skrzypczak and Mamcarz
2005a) and the position of stock releases, as a factor that contributes to occupying new
habitats by the pikeperch, kept decreasing together with the increase of the lake sizes
indicating migration of that species (Skrzypczak and Mamcarz 2005d).
The most of the attention was paid to the phenomena that took place in the exploited
populations of Coregoninae faced with intensive economical procedures carried out in the
worsening natural environment (Bnińska 2000). With the particular economic significance
of those species for the fishery management (Wołos 1998, Turkowski 2002), intensive
vendace and whitefish stock releases were carried out stating from the 1950s (Bernatow-
icz 1955, Szczerbowski 1977), which resulted not only in the increase of catches of those
species (Leopold et al. 1986) but also in the greater number of lakes that they occurred in
(Szczerbowski 1977, Skrzypczak and Mamcarz 2006b). Often, the vendace and whitefish
were moved to the lakes that generally did not fit into the conditions of their natural habi-
tats (Bernatowicz 1960, Szczerbowski 1969). This is confirmed by the results of the stud-
ied by Skrzypczak (2008) who noted the presence of Coregoninae in tench-pike type and
pikeperch type of lakes, proving as well their strongest connection with the vendace type
of lakes and a bit weaker one with the bream type of lakes. In the second half of 1960s,
an introduction of the third species of the Coregonus sp. type - the peled - was started in
Polish lakes (Bernatowicz and Sawicki 1971). As a part of those processes, hybridization
of the peled with the whitefish was made, which is deemed to be one of the causes of
the diminishing effectiveness in the stock management for those species (Mamcarz 1992).
With the deterioration of the environment, decreasing effectiveness of the Coregoninae’s
112
natural spawning was detected and presence of that fish in the lake ecosystems was to-
tally dependent on the stock releases (Leopoldet al. 1986). At the same time, together with
the advancing eutrophication, a decreasing effectiveness of fish stock releases was noticed
(Leopold and Wołos 1998). A breakdown in the Polish management of the Coregoninae
took place in the middle of 1980s (Wołos 1998).
Phenomena connected with the presence of the fish totally dependent on the stock
releases to the lakes were of quite different character. Of the most importance were
intensive eel stock releases, without which that species would have been of a marginal
significance in the structure of ichthyofauna of Polish lakes (Korulczyk 1978). For many
years, the eel played a pivotal role in the lake management. Therefore, a lot of attention
was paid to it in terms of estimation of intensiveness and effectiveness of stock releases
to various types of lakes and the efficiency obtained in them (Leopold 1974). The eel’s
share in the structure of the caught ichthyofauna did not depend directly on the wors-
ening state of natural environment resulting from advancing eutrophication (Bnińska
2000). With the growing catches of eel in the 1960s and 1970s, it was discovered that
the efficiency of that species kept growing indirectly in the lakes with the higher level of
their trophic status (Nowak 1978). The analysis of the eel share in the structure of the
caught ichthyofauna of north-eastern Poland in 1951 - 1994 showed unequivocally that
its highest percentage occurred in pikeperch type of lakes (Skrzypczak 2008).
In spite of criticism connected with introducing the carp into natural reservoirs, result-
ing from fears of its competition for food with the local fish and a lack of possibility to
obtain profitable production (Kulmatycki 1933), intensive stock releases of that species to
the lakes were started in 1951 (Nowak and Waluś 1969). In the light of assumed produc-
tion capabilities of the lakes in Poland, the carp raising was intended for 20,000 ha with
the planned natural capacity of 175 kg/ha (Rudnicki et al. 1971). It was assumed that with
release doses ranging to a few hundreds of young fish for 1 ha of lakes, no decrease of
catches of other fish was going to occur (Nowak and Waluś 1969). The revised number
of 635 lakes of north-eastern Poland in which presence of the carp was identified in the
structure of exploited ichthyofauna is an evidence of the range of the carp introduction
into Polish lakes (Skrzypczak 2008). Obviously, it must be assumed that intensive fish stock
releases within the frames of the carp management referred to a smaller number of lakes,
and the remaining cases of the carp occurrences in natural reservoirs were an effect of
unconsidered and causal fish stock releases and escapes of that fish from the breeding
facilities. The analysis of that species in the lakes of the north-eastern Poland indicates that
it was connected more strongly with the pikeperch type of lakes than with the tench-pike
type ones, as it occurred in a higher percentage of the exploited pikeperch lakes. At the
same time, the carp share in the total annual catch was greater in the tench-pike type of
lakes. It may be suspected that the pikeperch lakes were more favourable for the carp
when considering the requirements that set for the lakes chosen for the carp stock re-
leases – a theoretically less share of the pike in the structure of ichthyofauna and a smaller
number of water plants that made catches of that species more difficult.
The need for introducing into Polish lakes a species that would use plankton re-
sources was mentioned when starting the creation of a fishery management system
for the open waters (Stangenberg 1956). Faced with a rise of the trophic level of Polish
113
lakes in the late 1960, a procedure of introduction of herbivorous fish was started - the
grass carp, the silver carp and the bighead carp (Rudnicki et al. 1971). After several years
of protection, the exploitation of those species was started in early 1970. Despite great
expectations that the herbivorous fish would increase efficiency of the lakes by using
additional food base of eutrophic lakes, a range of difficulties connected with manag-
ing those species were shown (Krzywosz 1985). Studies by Skrzypczak (2008) indicate
that on the territory of the north-eastern Poland the most frequent occurrences of the
herbivorous fish in the structure of ichthyofauna were identified in the tench-pike type
(94) and bream type of lakes (85). At the same time, those species occur in the highest
percentage of the pikeperch type of lakes, which is in 48% of their total.
A wide panorama of changes in the formations of fish was presented by Leopold et
al. (1986) who included into his study an area of circa 95% of the surfaces of all the lakes
in Poland in 1956 - 1980. Within 25 years the authors noticed a constant increase in the
catches of the Cyprinidae fish (mostly silver bream, bream M and roach M, and in a lesser
degree bream D+S and roach S) as well as in the catches of a group of fish dependant on
stock releases that included: the carp, Coregoninae, the herbivore and the eel. A general
decrease in catches was identified for the littoral species (crucian carp, tench and pike) and
in a non-Cyprinidae group that (apart from the pike) included: the pikeperch, the perch
and the ruffle, as well as the smelt. The trend liners of those phenomena converged with
the theoretical eutrophication model described by Hartmann (1977) and suggested that
the Polish lakes at that time were between the 3rd and 4th phase of eutrophication. The
trends described by Leopold at al. (1986) on the basis of the changes in the catch volumes
manifest a character similar to the phenomena observed by Skrzypczak (2008). However,
a simulation mathematical model, applied by Leopold et al. (1986) to predict changes in
the structure of ichthyofauna in the next 15 years, which is to the late 1990, could not
include a change of trends that happened in the mid 1980s. After decreasing for many
years, a gradual increase of share in the structure of the exploited ichthyofauna, lasting
until the end of the analysed period, was noticed for the crucian carp, perch M, pikeperch,
pike, as well as for bream D+S (excluding the tench-pike type of lakes) and perch D+S (ex-
cluding the vendace type of lakes). At the same time, in the structure of the fish catches in
the lakes of the north-eastern Poland a decrease was noticed in the shares of the following
fish: the silver bream, roach S, whitefish and peled, as well as bream M (in the bream type
and vendace type of lakes), roach M (excluding the vendace type of lakes) and vendace
(in the bream type of lakes). A halt in the decreasing trend with no symptoms of increase
in the percentage was then characteristic for the tench, excluding the tench-pike type of
lakes in which the negative trend was observed until 1994 (Skrzypczak 2008).
Fishery typology versus eutrophication

The direction of changes in the lake ecosystems evoked by processes of eutrophica-
tion and resulting in the restructure of ichthyofauna was predicted by Stangenberg (1956).
Several years later, that phenomenon was confirmed by the studies by Colby et al. (1972),
Hartmann (1977) and Leopold et al. (1986). Stangenberg (19560 viewed the problem of
changes as the issue of a decline of the vendace and whitefish that have higher environ-
mental requirements, as well as in the phenomenon of over-stocking of perch-roach type
114
of lakes (according to the fishery characteristic of those times) with the pike and crucian
carp as the eutrophication increased. Simultaneously, according to that author, less and
less lakes would meet the requirements that were then created for the lakes of bream-
pikeperch character. Thus, it is very symptomatic that before finishing the works on the
fishery typology, a possibility of transition of one type of lake into another was assumed.
A partial confirmation of that thesis, in relation to the most shallow and the least morpho-
metrically differentiated lakes, was obtained by Skrzypczak (2008) in the phase of colleting
and preliminary analysis of the exploitation data form the lakes of the north-eastern Poland.
During the change of the patterns for the management books in the late 1960, a part of the
reservoirs classified in 1950s as the tench-pike type, after several years of exploitation, was
classified in new books as the crucian carp type. In many cases it means stopping mana-
gerial procedures in those lakes. At the same time, even if no changes were made to the
categorization, a gradual limitation of the exploitation and stocking processes took place in
the smallest reservoirs that usually belonged to the tench-pike type. A statistical significance
of the lowering index of the usage of the catch period, in a form of a visibly decreasing
number of months of exploitation in the lakes with the surface below 50 ha was identified
by Skrzypczak and Mamcarz (2003). A statistical decrease of the value of that parameter
was also noticed for the larger lakes but in a more limited scope. However, no case of a ty-
pological change was noticed for vendace type, bream type, or pikeperch type of lakes.
A number of observations regarding restructuring the fish formations were made
when studying the populations of vendace in Polish lakes. It was then that occurrences of
the pikeperch in new habitats, much different to the conditions of the pikeperch type of
lakes, was discovered (Wołos 1996, Skrzypczak and Mamcarz 2005a). The management
analysis for the Coregoninae resources made Bnińska and Wołos (1998) draw a conclu-
sion that the discussed predator is an index species for early eutrophication processes.
According to those authors’ suggestions, in an advancing process of eutrophication lakes
go through a so-called “pikeperch phase” when the population of that species increases
rapidly. Those observations may have only referred to the bream type and vendace type of
lakes that are the main habitats of the vendace. The occurrence of the “pikeperch phase”
most likely does not suggest a direct need for changes in classification of such lakes into
a different fishery type. However, it is difficult no to notice the fact that Bauch (1955) in his
classification of lakes singled out four types of pikeperch lakes, two of which he described
as relatively-deep lakes (15–30 metres). Considering the fact that eutrophication results in
lowered water clarity and oxygen shortages in bottom layers during the summer stagna-
tion (Leopold et al. 1986), types IIIa and IV of the pikeperch lakes according to Bauch’s
typology (1955) manifest properties of eutrophicated bream type and vendace type of
lakes. The Niegocin Lake may serve as an example here as its morphological properties
unequivocally indicated the vendace type. In the 1960s, the vendace was still exploited
there (Mamcarz and Skrzypczak 2002). At the same time, oxygen deficits were noticed at
the depth of several metres as well as the occurrences of hydrogen sulphide at the depth
of a dozen of metres (Zachwieja 1966). In the 1970s and 1980s, with no occurrences of
vendace in the structure of the exploited ichthyofauna, the Negocin Lake was one of the
main habitats of pikeperch in the complex of the Great Masurian Lakes (Skrzypczak and
Mamcarz 1995). Later on, regulations made to the management of water and sewages in
115
the drainage of the Niegocin Lake in the early 1990 within several years resulted in better
quality parameters of the lake waters and a change to the habitat conditions of the ichthy-
ofauna. It was then that a visible decline in the pikeperch population occurred (Skrzypczak
and Mamcarz 2005d), which is another proof of dynamic character of changes that take
place in lake ecosystems as well as an example of a specific revert of the trophic conditions
described by Gołębiewski (1976).
Fishery typology in the dynamic model

The analysis of a share if the fish formations (Coregoninae, Percidae, littoral ones,
and big and small Cyprinidae) in the structure of ichthyofauna in all the time phases
indicates a total difference of the tench-pike type of lakes when compared to the bream
type, pikeperch type and vendace type of lakes (Skrzypczak 2008). At the same time,
differentiated character of the environment in the tench-pike lakes provides the greatest
possibility of transforming it into the crucian carp type. According to a concept by Back-
iel and Zawisza (1956), the pikeperch type of lakes may be described as reservoirs with
incomplete differentiation of their environments, and the bream and vendace types may
be called complexly differentiated lakes. The greatest number of similarities between the
two latter types, as far their fish formations are concerned, were noticed in 1951–1964,
so in the lake environment that was theoretically least changed by the human. Against
the background of years-long anthropressure and environmental changes, a temporal
decrease of the similarity in the structure of the fish formations and an increase of the
similarity in the species structure of ichthyofauna are vey characteristic. Most probably,
it was visible when comparing bream type and vendace type of lakes. In 1981–1994, the
similarity of the exploited ichthyofauna between those two types referred to the pike,
crucian carp, roach and silver bream. It may be suspected that summer deficits in the
bottom layers, as a result of eutrophication, led to a significant assimilation of environ-
mental conditions in those stratified types of lakes. In some sense, the attribute that
differed them originally - the depth and with of the profundial accessible for fish – had
disappeared. This was to decide the most on the erasure of the differences between the
habitat conditions in the bream type and vendace type of lakes in 1981–1994 (Skrzyp-
czak 2008).
May it then be said that, in the context of the observed phenomena and with the as-
sumptions of the valid fishery classification, there are e.g. pikeperch type of lakes in the
basin of the Czarna Hańcza River if in the light of the studies by Skrzypczak (2008) they
were not present there a dozen years ago? Definitely not. However, it may be suspected
that some part of the bream type and vendace type of lakes in that territory, as well as
in the territories of other river basin, might be classified to one of the pikeperch lake
categories in the Bauch’s classification (1955), though a few years ago they did not meet
their requirements. It proves a static character of the fishery typology, valid for Polish
lakes. A change of the trophic classification by Neumann and Thienemann into a dy-
namic model, adjusted to the post-glacial reservoirs of Polish climate zone, was made
by Stangenberg (1936) and Wiszniewski (1953). What linked the natural succession of
lakes with the unavoidable succession of fish formations was the Colby curve (1972).
However, it was not until the dynamic model by Hartmann (1977) appeared that the
116
multidimensional process of changes in the lake environments evoked by their advanc-

ing eutrophication was considered.
The dynamic model of fishery typology must include the fact that the eutrophication
process generates groups of stratified lakes that in spite of morphological differences
manifest a significant similarity in terms of their changed temperature, oxygen, and habitat
conditions. This manifest also by a different level of water deoxygenation of the hypolimni-
on (in extreme cases also of the metalimnion) during the time of the summer stratification
and unavailability of the profundial. Such a function might be adopted by the category
β bream lakes, when assum-
ing a possibility of reverting
the properties in those lakes
based on the rule of the tro-
phism reversion (Fig. 1).
When observing the
pace and the effects of
changes in the lakes influ-
enced by eutrophication, it
Figure 1. Dynamic model for the fishery typology of lakes.
must be stressed that such
phenomena may occur in a time that may refer to the human life. However, it is difficult to
imagine how that limitation is passed by the processes of natural “ageing” and evolution
of lakes, which makes it possible for vendace type of lakes to transform into bream type
of lakes (category α) and to overcome the stratification barrier in a process of filling the
lake bowl with sediments or a process of getting shallow. Non-stratified lakes, that may
have properties of both pikeperch type and tench-pike type of lakes, are on the other side
of that barrier. When considering the morphological features of those reservoirs and the
so-far observed directions in the changes to their structure of ichthyofauna, a possibility
of transforming one type into another one should not be excluded. Biological studies
conducted in shallow eutrophic reservoirs, divided into “clear-water” macrophyte lakes
and “turbid-water” phytoplankton ones, provide a premise for such a thesis (Gąbka et
al. 2005, Iglikowska 2005). The depth index and the relative depth of the pikeperch type
and tench-pike type of lakes does not differ statistically, and occurrences of the pikeperch
in the tench-pike type of lakes is relatively frequent with an increasing trend (Skrzypczak
2008). It should be assumed that in the processes of getting shallow gradually and becom-
ing the land, which go beyond the possibility of observing it during a human life, those
lakes are to transform into the crucian carp type. Based on the long-term observations of
the phenomena taking place in a strongly eutrophic shallow lakes, it may be said that the
pace of transforming the pikeperch type and tench-pike type into the crucian carp type of
lakes may be accelerated by regular occurrences of winter oxygen shortages in them. They
dramatically reduce their structure of ichthyofauna to the state observed nowadays in the
crucian carp lakes.
Considerations on the changes in the structure of ichthyofauna that are to serve
as the basis for the lake classification should include the role of fish stock releases as
a modifier of fish formations. In Polish lakes, occurrence of eel and Coregoninae is totally
dependent on the stock releases and limitations to tench and pike stock releases would
117
most probably lead to a drastic decrease in the populations of those fish. The fishery
typology cannot ignore the state in which the leading species have lost their abilities
to exist in a for of natural populations in a numerous lakes and the occurrence of the
species in the lake ecosystem is charged with expenditures that are justified only in the
context of maintaining biodiversity. At the same time, the fishery classification must
make a difference between occurrences of a given species in a lake from the situation
when a lake make its habitat.
Summary
In the context of the above considerations, lobby discussions undermining the sense
of using the fishery typology for the effects induced by the lake eutrophication seem
groundless. In spite of time flow and the observed qualitative and quantitative changes
in fish formations, the diversification of the species structure of ichthyofauna in particu-
lar fishery types is still visible. Those phenomena were observed during fishery manage-
ment of Polish lakes, which included intensive fish stock releases and catches carried out
in the circumstances of non-market economy and against the background of changes
taking place in the environments of the lakes. The static character of the classification is
a limitation to the functionality of the fishery typology. Overcoming it is possible when
including the conception of a dynamic transformation of undisturbed and stratified ven-
dace type and bream type of lakes (α) into β bream type of lakes with various levels of
disturbance depending on their trophic phase and specification of their oxygen condi-
tions. Future observations of possible transformations of fish formations in Polish lakes
are necessitated by visible changes in the model of fishery management for the natural
water reservoirs, their increasing functions in tourism and recreation, as well as by oc-
currence of new modifiers. They may include climatic changes the results and scope of
which are unpredictable to a large degree.
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120
The state and prospects of the
fish management of coregonids
in the context of changes in the
fishery industry
Mirosław Szczepkowski1 Robert Stabiński2
1
Inland Fisheries Institute in Olsztyn, Department of Sturgeon Fish
Breeding in Pieczarki, szczepkowski@infish.com.pl
2
Polish Angling Association, Fish Farm in Suwałki,
pzw@suwalki.com.pl
Abstract
The purpose of this study has been to present the current state and prospects of pop-
ulations of coregonid fish in the context of the ongoing changes in the fishery industry.
There are two natives species of the genus Coregonus in Poland: vendace (Coregonus al-
bula (L.)) and European whitefish (Coregonus lavaretus (L.)), but two other species were
introduced in the 20th century: peled (Coregonus peled Gmelin) and muksun (Coregonus
muksun (Pallas)). Both of the native species of coregonids are among the most valuable
fish in fishery industry, but their present state is radically different.
Vendace, of which about 200 tons annually are caught, is in a far better situation. The
importance of this species in lake fisheries increases, largely due to declining catches of
a number of other valuable species, including eels (Anguilla anguilla (L.)). Populations of
vendace are strengthened by stocking, mainly in the form of mass introduction of hatch.
In 2009, over 340 million individuals of hatch were released to water bodies in Poland. Eu-
ropean whitefish has lost its position as an economically important species, and currently
attempts are made to preserve or recreate this species in selected lakes. For this purpose, cul-
ture spawning herds have been created in hatcheries, so that fish stocking material can be
produced annually. European whitefish, however, has a chance to revive because of its valu-
able contribution to aquaculture, especially in the facilities built to culture rainbow trout.
Introduction
There are two native species of coregonid fish living in Poland today. These are ven-
dace (Coregonus albula (L.) and European whitefish (Coregonus lavaretus (L.)). In the
20th century, two other species were introduced: peled (Coregonus peled Gmelin) and,
twice, muksun (Coregonus muksun Pallas) (Uryn 1987). These are, however, species for-
121
eign to our ichthyofauna (http://www.iop.krakow.pl/ias/) and now cannot be released to

Polish waters. Coregonid fish (Coregonus) are typical planktonophagues, which spend all
their lives in the open part of a lake, in the pelagic zone. Not many other fish dwell there,
which makes coregonid fish an important element in the biodiversity of our lakes.
Both coregonid species present in Poland are among the most valuable fish in the
fishery industry, the fact confirmed by the role they are assigned in fishery plans as well
as the value they attain when stockings are evaluated (Anonymous 2007, www.rzgw.
warszawa.pl). Although the two species are similar to each other in many respects (e.g.
habitat requirements, type of food, habitats), their present condition is radically differ-
ent. This paper contains a presentation of the current state of vendace and European
whitefish in lakes, especially in the Masurian and Suwałki Lake Districts.
Vendace
Vendace remains a very important species in lake fishery, and its catches are on
a high level, oscillating around 200 tons annually (Wołos et al. 2010).
It is justifiable to say that in the recent years vendace has become more important to
fishermen due to a decrease in catches of other valuable species, mainly eel Anguilla an-
guilla (L.). Another fact that raises interest in this fish is that vendace is not a game fish.
Catches of vendace are run from June to the end of September, mainly with gillnets,
and are characterized by a large variability over many years. This fact is of economic
importance because in the years when the amount of caught vendace is high, its price
declines. In some years, it was even necessary to discontinue catches. Another significant
factor is the size of captured fish. The most desirable are fish which weigh over 100 g, as
these are sold at the highest prices. Fish weighing 50-60 g each are sold much cheaper,
down to 50% of the maximum price. Besides, when the fish are small, they are difficult
to market and therefore their catches sometimes have to be limited.
Because it is necessary to obtain fish stock material, a large proportion of catches
falls on the spawning season. It is estimated that during the spawning, about 20% of
the annual catches are done. At that time, catches are mainly carried out in lakes where
strong populations of vendace appear. A negative aspect of catching the fish during
the spawning season is the fact that fish after spawning are less valuable for processing
(especially females), which is why their price falls down by about 30%.
The effects of vendace stocking are evident, which is why it is one of the most impor-
tant species in the fish management. Production of stocking material is mainly based on
obtaining wild spawners and conducting their artificial reproduction (Fig. 1). For many
years, the primary form of fish stock is hatch. In 2009, over 340 million individuals of
fish hatch and only about 4 million of summer fish fry were released (Mickiewicz 2010).
Attempts have been made to rear the hatch to older stages, up to the formation of
spawning herds (Wziątek et al. 2009). However, it seems likely that in the nearest future
the present model of stocking, based on large amounts of hatch, will dominate.
European whitefish
In the past, European whitefish was a species of a considerable economic importance
and its catches reached 100 tons a year (Szczerbowski 1993). In the 1970s, the catches
122
The state and prospects of the fish management of coregonids in the context of changes…
of this fish in some single

lakes were larger than the
present catches for all Po-
land (Falkowski 2004).
There are many forms
distinguishable among Eu-
ropean whitefish, which
differ from one another in
the conformation, shape of
the mouth as well as some
meristic features, among
which the number of gill-
rakers is the most impor-
tant in respect of this spe-
cies. The European white-
fish inhabiting our lakes Figure 1. Incubation of eyed vendace eggs at the hatchery
of the Polish Angling Association in Doliwy.
are classified as Coregonus
lavaretus generosus, although in some lakes local populations have been distinguished,
for example in Miedwie and Wigry Lakes (Lityński 1932) and some researchers (Sobieska
and Gecys 2007) now claim that the European whitefish in Wisztynieckie Lake is a sep-
arate population (houting whitefish Coregonus lavaretus oxyrhynchus), which is also
used for stocking lakes in Masuria (Mazury).
European whitefish belongs to the species which have long been supported by fish-
ermen so that artificial reproduction and incubation of eggs of this fish started in the
early days of the hatchery production (Benecke 1881). This, however, was also the begin-
ning of hybridization of populations of whitefish from different water bodies and even
from different geographical areas. Let us take as an example the stocking described by
Benecke (1881), when European whitefish from the Curonian Lagoon was released to
Narie Lake. Today, we know that this was most probably migratory whitefish. This proc-
ess of crossing fish populations often was completely uncontrollable, therefore it seems
justifiable to treat the European whitefish in Poland as intermediate varieties, which
some authors have been suggesting for a long time (Terlecki and Kempińska 1956).
Unlike vendace, the abundance of European whitefish has been much more depend-
ent on human activities. In the past, there were a few periods of time when catches of
this species increased considerably after intensive fish stocking, for example in the 19th
century or after World War Two. At present, the situation is much more difficult because
of the enormous changes that have taken place in the lakes and in their immediate sur-
roundings. We mean water pollution, causing oxygen deficit, reduced area of parts of
lakes available to fish, muddy spawning grounds and big populations of black cormo-
rant. Moreover, the state of European whitefish populations has been adversely affected
by hybridization with other fish of the genus Coregonus, mainly with peled (Łuczyński et
al. 1992). Unfortunately, milt of vendace is still used for fertilization of European white-
fish spawn (own observations).
Some influence on the state of the resources of European whitefish has also been
produced by the change which has occurred in the way lakes are managed now, that
123
is limited catches of eel and

more importance attached
to the European whitefish
fishing. Abandoning the
catches with dragged tools
in the summer and replac-
ing them with gillnets has
had a negative influence
on European whitefish by
partly eliminating their ju-
venile specimens. Unlike
vendace, European white-
fish is now fished mainly
during the spawning sea-
Figure 2. European whitefish captured in Mamry Lake.
son (Fig. 2). Captured fish
are transported live to hatcheries where they are spawned. Fishermen receive additional
income for the spawn. In other seasons of the year, European whitefish is caught in
these lakes where it has been intensively stocked and which do not provide conditions
for its natural reproduction (e.g. Nidzkie Lake).
Although European whitefish has lost its economic importance, it is still valued by fish-
ermen, which is mainly a consequence of their being used to this species. This is the reason
why some actions have been undertaken to ensure a safer future for the species, such as:
• creating spawning herds under controlled conditions
• obtaining spawn from cultured spawners
• producing hatch and fry for releases to selected water bodies
• obtaining (as much as possible) wild spawners in order to sustain genetic variation
of the population of European whitefish.
Following genetic studies, two European whitefish spawning herds are now main-
tained, originating from Gaładuś Lake and from Wisztynieckie Lake (Szczepkowski et al.
2010). These herds are kept in two fish breeding stations: the Department of Sturgeon
Fish Breeding in Pieczarki (the Inland Fisheries Institute) and the Polish Angling Associa-
tion Fish Hatchery in Gawrychruda (Szczepkowski et al. 2008). In the recent years, about
70-80 liters of spawn have been obtained annually. This is enough to produce the stock
material for fish releases and any excess spawn which cannot be taken for further rear-
ing is used for replenishing European whitefish populations in many fish farms.
The results of fish stocking suggest their low effectiveness, especially when hatch
and the youngest juvenile stages are used. Better effects are achieved when older forms
of the fish stock are taken, especially autumn fish fry. This is most probably related to
their higher tolerance to predators, particularly perch (Perca fluviatilis (L.)). However, it
is not easy to rear adequate amounts of autumn fish fry. In water recirculation systems,
the two problems are: limited capacity of tanks and considerable costs of rearing older
material. In ponds, in turn, there could be some problems in summer due to high tem-
peratures and oxygen deficits. In earthen ponds, catching reared fish fry is very difficult
124
The state and prospects of the fish management of coregonids in the context of changes…
and is sometimes associated with large losses. Much better effects are obtained when
whitefish fry is reared in trout ponds. It has even been tried to rear European whitefish
to marketable size, e.g. in the Fish Farm in Ruś near Olsztyn. Such attempts should be
encouraged for the sake of preserving the species, provided that the initial material
originates from genetically pure populations, as is the case in the Fish Farm in Ruś. Then,
the reared material can be used for stocking and be treated as a reserve resource for the
maintained herds of spawners. The pond culture experiments run for a few years now
enable us to conclude that the biggest threat to European whitefish is its sensitivity to
manipulations and diseases, particularly bacterial ones. European whitefish, however,
seems more tolerant to the environmental conditions (temperature, oxygen concentra-
tion, amount of suspended matter) than has been expected.
Summary
Vendace and European whitefish now have a completely different position in the
fishery industry. A good example confirming this statement is the volume of catches of
spawners during the protection period, which is several-fold higher for vendace (Zakęś
and Danilewicz 2010).
It seems that in the nearest future vendace will not be endangered and its impor-
tance in the lake economy might even grow. In contrast, European whitefish will not be
able to regain high abundance of its populations in the predictable time. It is, however,
possible to recreate and maintain populations of this species in some water bodies.
Thus, in order to attain superior results, it is recommendable to focus on the several sites
where the effects of stocking are the best.
The status of lake fish farms will be of a large importance to the future of European
whitefish. If their economic situation continues to deteriorate, European whitefish may
suffer because then it can be expected that fish farm managers will focus on the spe-
cies which yield the best effects. Unfortunately, European whitefish does not belong to
such species. Therefore, for the preservation of this species in lakes, the lake owner (the
State Treasury represented by the Regional Water Managements Boards) should prepare
mechanisms which will encourage fish farms to continue European whitefish stocking
even at low effectiveness.
Extremely important for the protection of European whitefish are the activities which
involve creating herds of spawners in hatcheries and improving the effectiveness of
hatchery procedures and the biological quality of the stocking material. Aquaculture
seems to bring some hope for the future of this species, especially when conducted in
the facilities where rainbow trout (Oncorhynchus mykis Walbaum) is cultured. The expe-
rience gained from the work done in Ruś shows that European whitefish is much more
tolerant to high temperatures than rainbow trout. This is also confirmed by the observa-
tions of the breeding of spawners, carried out for many years now at the Department of
Sturgeon Fish Breeding. Normal feeding of European whitefish there takes place at the
temperature of water temperature reaching 24oC. Owing to this characteristic, European
whitefish can become a valuable species supplementing production in these fish culture
facilities, where there is a risk that summer temperatures may rise so as to endanger
rainbow trout culture.
125
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126
Effect of northern pike Esox
lucius L. on a spawning
population of vendace
Coregonus albula (L.)
in Wigry Lake
Andrzej Martyniak, Katarzyna Mierzejewska,
Katarzyna Stańczak, Michał Osewski1,
Kamil Kisielewski, Michał Juńczyk,
Łukasz Dziemian, Paweł Zawal
Department of Fish Biology and Breeding, Faculty of Environmen-

tal Protection and Fisheries, University of Warmia and Mazury in
Olsztyn, kbhr@uwm.edu.pl
1
Wigierski National Park in Krzywe
Abstract
In four consecutive years (from 2005 to 2008), northern pike was caught from the
sublittoral zone in Wigry Lake during spawning season of vendace. Analysis of stomach
content of 243 predators, has shown that the population of vendace in Wigry Lake is
subjected to strong predation pressure of northern pike, with contribution by count in
the diet from 84.9 to 96.8%. Roach and perch represented (respectively) 8.6 and 9.6% of
the northern pike’s prey. There was no evidence for the predator-prey size relationships
because the size of prey fish was conditioned by the size of vendace ready to spawn. In
the analyzed population it was confirmed that the availability of prey decidedly influ-
enced the pike’s food selection, individuals of the length over 38 cm (SL) expanded their
feeding zone beyond the littoral and the negligible role of cannibalism in the studied
population. The relieved predatory pressure may be associated with the fact that north-
ern pike preferred individuals which had already spawned.
Introduction
Northern pike is a widespread fish in Polish waters. Adults are obligatory predators,
choosing various species of fish-prey according to its availability, that is density, distribu-
127
tion, daily and seasonal activity, the fish condition and health as well as the presence
of effective and numerous hiding places. In lakes in Central Europe, northern pike most
often catches roach and perch (Antosiak 1963, Załachowski 1965 and 1973); in rivers
it also attacks salmonid fish. When food is sufficiently abundant, cases of cannibalism
do not exceed 1-2% (Frost 1954, Lawler 1965). Other aquatic vertebrates appear in the
diet rarely, in small numbers and rather periodically. The food selection of this predatory
fish is largely dependent on its ‘being used to’ a certain species of prey, that is on spe-
cific trophic relations formed and strengthened under the conditions provided by a given
aquatic ecosystem. Pike can regulate populations growth of fish co-occurring in water
body, by this way it is able to shape quantitative ratios in the fish community and limit
the effectiveness of stocking with valuable species. Vendace (Coregonus albula L.) is a relic
species, one of the most valuable fish in the Polish ichthyofauna in terms of the wildlife
and economy. In 1951-1994, it was fished from 463 lakes in north-eastern Poland. The
most dynamic growth in the commercial exploitation of this species was recorded in the
1950s (Skrzypczak 2008). At present, due to the accelerated eutrophication of lakes, the
living and reproduction conditions of vendace have become considerably inferior. Vendace
has disappeared from many Polish lakes as a result of very low or zero effectiveness of
natural spawning. In order to sustain the populations of this species, fish stock material
produced in hatcheries is released to many water bodies. It is presumed that the most
numerous population of vendace in Poland dwells in Wigry Lake; its estimated biomass
is 222 kg/ha (Świerzowski 1999). Wigry Lake provides 10% of the vendace captured in
Poland (in 1979-2004, about 6 kg/ha). After World War One, most probably due to the
badly run fishery management, the population of vendace in Wigry Lake almost disap-
peared (Lityński 1924a, Lityński 1924b). Successful reintroduction of this species required
intensive fish stock releases, which have been regularly conducted in Wigry Lake since the
1930s (Białokoz et al. 1999). With the lake’s bottom becoming muddy and some local
oxygen deficits, the success of natural spawning of vendace is very limited. In the genera-
tions 2000 and 2001, 80 and 60% of individuals, respectively, originated from a hatchery
(Kozłowski 2007). Restitution of lake trout and supporting the population of northern pike
in Wigry Lake likewise require regular fish stocking (Białokoz et al. 1999).
The co-occurrence of northern pike and vendace in Wigry Lake raises questions regard-
ing the predatory pressure on the population of this valuable fish. Vendace feeds on zoo-
plankton and is a pelagic gregarious fish, which migrates in search of food and spawning
grounds. For northern pike, the natural zone for life and hunting, especially in the early
life, is the littoral. Older individuals (particularly above 45 cm in body length) expand their
foraging area over deeper layers of water, such as sublittoral, profundal and even pelagial
zones (Dziekońska 1954). It is highly likely that the two fish species will meet, especially in
the sublittoral zone, where vendace gathers for spawning, forming large stocks and thus
becoming a source of food, easily available and valuable to larger pikes.
Wigry Lake has a specific, unique character, mainly because of the varied glacial land-
scape surrounding the lake. As such, it has long been considered as a natural treasure.
The first concept of creating a nature reserve that would encompass Wigry Lake ap-
peared in the 1920s (Hryiewicz and Lityński 1942a, Lityński 1924b). In 1975, Wigry Lake
was registered as the most valuable water bodies in the world, and in 1989 Wigierski Na-
128
Wpływ populacji szczupaka (Esox lucius L.) na populację tarłową sielawy…
tional Park was established,

one of the largest in Poland
as it covers 15,086 ha, in-
cluding 2,908 ha of water
bodies, of which nearly
2,187 ha is the area of
Wigry Lake (Fig. 1) (Czeczu-
ga and Gołębiewski 1976).
The lake has been classi-
fied as meso-eutrophic and
a vendace-type lake (Szcz-
erbowski 1981, Brzęk 1988,
Kamińska 1999).
Protective fishery man-
agement in the lake com-
prises the regular and con-
trolled catches as well as
regular fish stocking. The
Figure 1. Bathymetric map of Wigry Lake
main aim of catches is to
reduce the excessively growing populations of small cyprinids and, once a year, com-
mercial catches of growing cohort of vendace. Pike is a species desirable and favored in
the ecosystem, serving to select and regulate abundance of co-habiting fish species, but
is subjected to strong man-made pressure even though its catches are regularly limited
(Świerzowski 1999, Kozłowski 2007).
Material and methods

Food composition of northern pike in Wigry Lake has been analyzed on the basis of
the stomach contents of 243 individuals captured in November 2005 and November
2006, from September to December 2007 and from September and December 2008
(Tab. 1). The fish were caught with gill nets of the mesh size 22x26 mm, killed immedi-
ately after catching and deep frozen. In a laboratory, defrosted fish were measured up
to 0.1 cm; the standard length (SL) and weight were used in further analysis. Afterwards,
stomachs of all the fish were dissected. In order to assess the foraging intensity of the
predator during the analyzed period, the percentage of full and empty stomachs was
calculated (Tab. 1).
The next steps involved the qualitative and quantitative analysis of the gastric con-
tents. The prey fish were species identified according to the external taxonomic features,
largely digested specimens – on the basis of osseous elements, otholits, scales, bones
and, in the case of cyprinids, the pharyngeals.
The length of partly digested fish was calculated using the following conversion fac-
tors (acc. to Horoszewicz 1964):
for roach SL = 12.9 x h + 0.23
where:
129
SL - the fish’s standard length (in cm)

h – height of the pharyngeal arch (in mm)
The length of slightly digested vendace was measured directly, the length of more evi-
dently digested ones, was assessed on the basis of the proportions of remaining body parts
according to the biometric features determined for the population (Kozłowski 2007).
The weight of the prey fish was calculated from the length (L) - weight (W) relation-
ship according to the commonly used formula W = k x Ln, where k and n are coefficients
characteristic for a population. In Wigry Lake, for roach k = 0.0141 and n = 3.1994 (Ter-
lecki 1987); for vendace k = 0.0102 and n = 3.0635 (Kozłowski 2007). The identification
of sex and gonadal maturity of prey fish were done whenever possible. The importance
of particular food components in the diet was assessed using the methods gathered by
Hyslop (1980): the contribution by number, by weight (using direct weighing or calcu-
lated from L-W relationship) and frequency of occurrence (percentage of stomachs filled
with particular item). The predator-pry size relationship was analyzed by the Pearson’s
correlation coefficient (at the significance level α=0.05).
Table 1. Standard length (SL) and percentage of foraging and non-foraging northern
pikes in samples from Wigry Lake.
Number Body length SL (cm) Full stomachs Empty stomachs

Date of catches
of fish average (min.–max .) (indiv.) (%) (indiv.) (%)
Nov. 2005 31 48.6 (43.9 – 57.2) 21 67.7 10 32.3
Nov. 2006 57 49.6 (38.0 – 62.0) 37 64.9 20 35.1
Sept./Dec. 2007 100 47.2 (38.0 – 68.0) 47 47.0 53 53.0
Sept /Dec. 2008 55 47.4 (39.5 – 66.0) 31 56.4 24 43.6
In total 243 48.2 (38.0 – 68.0) 136 59.0 107 41.0
Results
During the sampling seasons, the percentage of foraging northern pikes ranged from
47.00 to 67.74% depending on the year of study (Tab. 1).
The composition of northern pike’s food

In the stomachs analyzed from 2005 to 2008, 7 fish species in total were determined.
Vendace appeared most frequently and in the highest number (up to 8 individuals in one
stomach); smaller contribution belonged to roach and perch. There were just single in-
dividuals of northern pike, European whitefish, rudd and three-spined stickleback. Apart
from the fish, one individual of the crayfish Orconectes limosus was found (Tab. 3).
In the 2005 sample, vendace dominated both in the number (85.7%), weight (76.7%)
and frequency of occurrence (85.7%). The respective percentages for roach were 8.5, 7.7
and 9.5%. Besides, there was one three-spined stickleback and one northern pike (30 cm
l.c. in a cannibalistic northern pike of 53.5 cm l.c.) found (Tab. 3).
In the 2006 sample, two species were found, vendace prevailed, with the contribution
by number and by weight over 95% and the frequency of occurrence about 94%. Roach
was found with frequency 8%, contribution by number 3.2%, by weight 3.7% (Tab. 2).
130
Table 2. Contribution by number (CN), contribution by weight (CW) and frequency of occurrence (FO) of prey in the diet
of northern pike from Wigry Lake, determined in the autumn and winter season in subsequent years of the study.
Year 2005 2006 2007 2008

Prey CN (%) CW (%) FO (%) CN (%) CW (%) FO (%) CN (%) CW (%) FO (%) CN (%) CW (%) FO (%)
vendace 85.7 76.7 85.7 96.8 96.3 94.4 86.9 85.1 82.9 84.9 85.0 78.6
European
- - - - - - 0.9 5.7 2.1 - - -
whitefish
roach 8.6 7.7 9.5 3.2 3.6 8.3 7.5 6.2 8.5 4.1 10.0 10.7
perch - - - - - - 3.8 2.6 4.4 9.6 4.0 7.1
rudd - - - - - - - - - 1.4 1.0 3.6
pike 2.86 15.0 4.8 - - - - - - - - -

three-
spined
2.86 0.5 4.8 - - - - - - - - -
stickle-
back
crayfish - - - - - - 0.9 0.4 2.1 - - -
In total 100.0 100.0 104.8 100.0 100.0 102.8 100.0 100.0 48.0 100.0 100.0 100.0
Table 3. Number and size of the prey of northern pike from Wigry Lake in subsequent years of the study.
2005
N Standard length SL (cm) Body weigh (g)
Prey N Np
max. Range Average Range Average
vendace 30 1.7 4 13.0 – 17.2 15.00 26.4 – 62.2 41.20
roach 3 1 2 11.2 – 11.8 11.60 32.1 – 38.0 36.03
northern pike 1 1 1 30.0 30.00 _ _
three-spined
1 1 1 _ _ _ _
stickleback
2006
vendace 90 2.5 8 10.0 – 18.0 14.30 11.8 – 71.5 38.00
roach 3 1 1 11.7 – 12.5 12.10 36.9 – 45.6 41.60
2007
vendace 93 2.5 8 8.0-20.0 14.55 8.0-120.4 24.73
perch 8 2.6 5 6.5-14.5 8.19 8.1-17.9 8.34
roach 4 1 1 7.5-17.0 12.80 15.5-100 36.00
European
1 1 1 18.0 18.00 110.0 110.00
whitefish
crayfish 1 1 1 8. 5 8.50 20.0 20.00
2008
vendace 62 3.5 8 12.0-20.0 15.56 8.2-34.7 28.23
perch 7 3.5 5 7.1-14.8 11.03 9.0-19.7 13.13
roach 3 1 1 8.9-19.3 13.80 17.6-114.0 41.30
rudd 1 1 1 12.9 12.90 15.4 15.40
N – total number of prey fish in a sample
NP – average number of prey fish in a stomach
Nmax – maximum number of prey fish in a stomach
( _) – no data
131
In the sample obtained in 2007 (in September, November and December in total),
vendace was also the most important food item with contribution over 85% and fre-
quency, 82.9%. At the same time, roach made up 7.5% of prey fish. The share of perch
did not exceed 3.8% of ingested prey. Moreover, one European whitefish and one cray-
fish Orconectes limosus were found (Tab. 2). In the particular months of catches, the
percentage of vendace by number, by weight and frequency of occurrence were, respec-
tively: 80.0%, 82.8% and 66.6% in September, 86.0%, 84.6% and 89.4% in November
and 93.3%, 95.8% and 83.3% in December.
The diet of northern pike in 2008 (in September, November and December in total)
consisted of four components. The most numerous was also vendace, whose quanti-
tative and weight shares were about 85%, whereas the frequency of occurrence was
78.6%. The other fish were clearly less important in the diet (Tab. 2).
Size and number of the northern pike’s prey
In the food of the fish analyzed in 2005-2008, whose body length ranged from 38
to 68 cm (l.c.) (Tab. 3), 307 fish in total were found, including 275 vendace, 15 perch
and 13 roach, at their maximum number (respectively) of 8, 5 and 2 individuals in one
stomach (Tab. 3). Besides, single individuals of rudd, European whitefish, northern pike
and three-spined stickleback were found. The standard length (SL) of the prey fish (all
species) was from 6.5 cm (perch) to 20 cm (vendace), at the weight of 8.0 g (vendace)
to 120 g (vendace) (Tab. 3).
The mean standard length of ingested vendace in consecutive years of study was
quite similar and within the range of 14.30 to 15.56 cm (Tab. 3). The length of eaten
roach was within 7.5 and 19.3 cm, and perch – from 6.5 to 14.8 cm (Tab. 3).
No significant correlation between the body length of the predator and the body
length of its prey, including vendace, was found. Person’s correlation coefficients in the
consecutive years of the study reached the values r<0.25 and were statistically non-sig-
nificant (p>0.05).
The sex ratio in 2007 sample (36 vendace) was determined. The fish comprised 11
females (30.5%) and 25 males (69.5%). Gonadal maturity of just 16 fish was possible to
evaluate; they were three females before spawning, three spawned females, one male
before spawning and 9 spawned ones.
Discussion
In many publications on the foraging of northern pike in Polish lakes, the authors claim
that roach is the main food item for this predator, with perch taking the second place
(Dziekońska 1954, Antosiak 1963, Białokoz and Krzywosz 1976, Stec 1980, Przystawik 1994,
Załachowski 1965). In rivers (the Vistula River, the basin of the Łyna River), white bream,
common nase and common dace, apart from roach, fell prey to pike (Horoszewicz 1964,
Martyniak et al. 1976). In the Cam River (Eastern England), 42% of components was three-
spined stickleback, with gudgeon (21%) being another important food item (Hartley 1948).
Variety of factors, such as the abundance, density and morphology of prey fish, may modify
the diet of pike, provided that the predator and its prey occupy the same ecological niche
(Dziekońska 1954, Antosiak 1963, Horoszewicz 1964, Załachowski 1973). Many authors
state that the qualitative and quantitative composition of northern pike’s food reflects the
132
fish community structure in a given water body (Dziekońska 1954, Antosiak 1963, Białokoz
and Krzywosz 1976). The weight contribution of particular fish species in the northern pike’s
food often diverges from the weight share recorded during commercial catches. There are
two reasons. On the one hand, there is a diversity of habitats occupied by different fish
species. On the other hand, commercial catches are selective and neglect less valuable fish
species. When control catches had been carried out using a drag nets (of a small mesh size)
within the zone inhabited by northern pike (in the littoral), the weight contribution of prey-
fish in stomachs of the northern pike was similar as in the catches. Our study conducted
in Wigry Lake supports the hypothesis that the availability of prey is the key to the choice
made by northern pike when hunting. Pikes measuring over 38.0 cm of standard length
(about 45 cm of total length), able to expand their foraging grounds beyond the littoral,
and captured in the areas where vendace gathered for spawning, mostly ate this fish (from
78.6 to 94.4% in particular years). Over the four years, in the autumn and winter seasons,
vendace constituted up to 96.3% of the weight of northern pike’s prey. One predator was
able to catch up to 8 individuals (Tab. 3). The season of reproduction is the time when fish
are more vulnerable to predatory pressure because they gather into spawning grounds (An-
tosiak 1963). The schooling behaviour is typical for vendace, therefore its populations could
be exposed to reinforced predatory pressure. Our study has confirmed that the population
of vendace in Wigry Lake faces such pressure exerted by northern pike every year. In other
lakes, too, mass foraging of vendace during its spawning season by pikes has been noticed
(Dziekońska 1954). Other fish species during their reproduction become a dominant compo-
nent in the predator’s diet. In Dobskie Lake, in May, stomachs contained only roach, but in
summer, when bleak spawns, it was detected in food of 88.2% of predators, while its aver-
age frequency of occurrence covering different seasons was 10.7% (Dziekońska 1954). The
northern pike’s pressure on vendace in Wigry Lake became evident even before the spawn-
ing season. In September 2007, vendace occurred with frequency 66.6% and prevailed in
the diet with contribution 80%.
The size of prey fish in the subsequent years was quite similar. The northern pike’s
diet was dominated by vendace ready to spawn, i.e. fish of a certain size. Therefore,
the predator-prey size relationship was not evident. Identification of sex and gonadal
maturity was done in sixteen vendace ingested. This group of fish was dominated by
spawned males. It is difficult to make a firm statement, based on the above result, that
the predatory pressure had been relieved. Further research would be necessary to verify
such a tendency. Nonetheless, the fish weakened by spawning are naturally more vulner-
able to a successful attack of a predator.
The second and third place in the northern pike’s diet were occupied by roach and perch
(Tab. 2), which are very popular in Wigry Lake. There was only one registered case of can-
nibalism (Tab. 3). Negligible role of cannibalism in northern pike’s diet in Wigry Lake may
be connected with the abundance of alternative prey species and with the relatively small
density of the pike population, despite of its regular stocking and limitation of catches.
Conclusions
1. The population of vendace in Wigry Lake during spawning and immediately before
it is subjected to strong predatory pressure exerted by northern pike.
133
2. During our study, the predator-prey relationship was not found because the size of
northern pike’s prey depended on the size of vendace entering the spawning season.
3. It was confirmed that the availability of prey played the predominant role in the
northern pike’s food selection, expansion of the foraging zone by northern pikes over 38
cm long (SL) beyond the littoral and negligible importance of cannibalism in the studied
population.
4. The results suggested that some predatory pressure of northern pike on the popu-
lation of vendace could be relieved by the predator focusing on spawned individuals
(specially males). Due to a small number of vendance classified in terms of gonadal ma-
turity, this observation needs to be verified.
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135
Formation of the ichthyofauna
in the Czorsztyński Dammed
Reservoir (the Dunajec River
system – southern Poland)
Leszek Augustyn
Polish Angling Association, Nowy Sącz Region,

33-300 Nowy Sącz, ul. Inwalidów Wojennych 14,
e-mail: leszek.augustyn@pzwns.info.pl
Abstract
The Dunajec River is the second largest tributary of the Vistula River in the Carpathian
Mountains. In 1997, at the border of two regions, Podhale and the Pieniny Mountains,
two reservoirs were built on that river: a large dammed reservoir called the Czorsztyński
Reservoir (1,266 ha) and a small equalization one in Sromowce Wyżne (88 ha). In the
1960s, the presence of 12 fish species was documented in the area planned to be turned
into the reservoir’s bowl in the future. The fish were dominated by rheophilic cyprinid
fish: nase (Chondrostoma nasus, 30.6%), barbel (Barbus barbus, 26%) and spotted bar-
bel (Barbus carpathicus, 11%). When the construction work on the dam and the reser-
voirs became more intensive (1981-1996), river gravel in the reservoir bowl was heavily
exploited, as a result of which several gravel pits appeared, in which the succession of
fish ran a similar course as in dammed reservoirs, i.e. from the dominance of roach and
pike, to superdominance of common bream. The ichthyofauna from these water bodies
spread to the Dunajec River, gradually supplanting rheophilic species, a situation that
was additionally compounded by the increasing pollution of waters with household and
industrial wastewater and sewage.
In 1992, a fishing and angling plan was accepted for the management and exploi-
tation of the Czorsztyński Reservoir. Salmonid species, such as lake trout (Salmo trutta
lacustris) and huchen (Hucho hucho) were taken as dominant predators. The fact that
fish were spread from the post-exploitation gravel pits into the reservoir being filled up
with water put at risk the whole concept of running the fishery economy with lake trout
and huchen as the key predators. In 1995, over 24,000 fish of the total weight over 8
tons were captured from these reservoirs by fishermen. Pike Esox lucius regulating the
number of fish and thus keeping the whole fish assemblage in a state of relative equi-
137
librium, lost any protection. It is estimated that in 1998, from the ice cover thaw to the
end of the year, over 6 tons of pike were caught, mainly spawners entering the Dunajec
River in search of spawning grounds.
Until 2002, the Czorsztyński Reservoir had been stocked with 1.89 million individuals
of larvae and 13 thousand individuals of fish fry of huchen as well as 270 thousand indi-
viduals of larvae and 13 thousand individuals of smolt of lake trout. The fact that these
fish releases had no evident success stimulated a change in the concept of the future
exploitation of the reservoir. It was assumed that pike, pike-perch (Sander lucioperca)
and, later, catfish (Silurus glanis) would keep a certain degree of predatory pressure,
necessary to maintain a proper dynamic equilibrium in the fish assemblage. Until the
end of 2008, the reservoir had been stocked with 654 thousand individuals of fry and
132 kg of spawners of pikeperch, as well as 3 thousand individuals of summer fry and
3.5 tons of autumn fry of pike.
The excessively abundant catches of pike carried out in 1997-1999 caused a collapse
in the structure of the ichthyofauna and that predator practically disappeared from the
lake. The fact that the trophic chain was therefore shortened, caused a specific popula-
tion explosion of roach, previously limited by the pike’s predatory pressure. In 2000-
2001, roach made up 58% qualitatively and 34% of the biomass of the ichthyofauna in
the Czorsztyński Reservoir. At present, its angling catches exceed 6.5 tons annually. Since
2003, owing to fish releases, the population of pike has been partly rebuilt. Next to pike-
perch, pike is now the main component of game catches of predatory fish. In the early
stage of building the ichthyofauna in the Czorsztyński Reservoir, common bream was
the most dynamic species. Its older generations, originating from the spawning events
taking place in the gravel pits dominated game catches. However, since 2003, catches
of bream have been falling and the individuals caught tend to be smaller. Changes in
the catches of bream, a species used as an indicator of lake eutrophication, suggest that
the increase in the trophic state of the reservoir has been halted. As the recent studies
demonstrate, due to some transformations in the land use structure in Podhale, nitrogen
loads from large-area pollution in the Dunajec River have decreased 11-fold over the
past 15 years. Is it therefore possible to return to the concept of stocking the reservoir
with fish fry of huchen and lake trout? Certainly not! The formation of the ichthyofauna
in the Czorsztyński Reservoir runs according to the course that is known from other
artificial lakes. In this case, the imperfection of the guidelines suggested by science has
been painfully verified by nature.
Introduction
Construction and operation of dams cause much damage to communities of aquatic
organisms. By dividing into two parts a water system that previously was a uniform en-
tity, fundamental changes in the structure and functions of live elements which inhabit
this system are stimulated (Penczak 1994). Dams destroy what is crucial to any river
– its continuum (Penczak et al. 1998). Dammed reservoirs created by raising rivers are
aquatic ecosystems that are different from stagnant waters or rivers, which is a conse-
quence of the specific character of transformations such reservoirs undergo (Starmach
1988). Research on ichthyofauna in artificial lakes pays much attention to modifica-
138
Formation of the ichthyofauna in the Czorsztyński Dammed Reservoir…
tions in structures related to the formation of a whole fish community (Starmach 1986,
Jelonek and Godlewska 2000, Gido et al. 2000, Matthews et al. 2004). A newly created
reservoir is an environment in the process of being shaped. Soon after it is filled up with
water, a specific natural race starts in order to colonize it (Frankiewicz 2000, Augustyn et
al. 2003). The initial development of a new fish assemblage depends on the state of the
ichthyofauna in the river before it was dammed (Mastyński 1985). In time, typical river-
ine fish species recede and the ones more adaptable to new conditions succeed (Jelonek
and Amirowicz 1987). The latter include the most expansive populations, such as roach
and common bream (Bieniarz et al. 1990).
The purpose of this study has been to characterize qualitative and quantitative
changes in the structure of the ichthyofauna in the Dunajec River since the construction
of the Czorsztyński Reservoir started, and the formation of the fish community after the
reservoir had been filled up with water. The characterization has been based on results
of research reported by many authors.
The hydrotechnical constructions on the Dunajec River

The Dunajec River is the
second largest Carpathian
tributary river of the Vis-
tula River. Its springs are in
the Tatra Mountains, from
which it flows through the
Orawsko-Nowotarska Val-
ley, cuts across the Pieniny
and the Beskidy Moun-
tains and then flows into
the Sądecka Valley, where
it receives its largest tribu-
tary, the Poprad River (Fig.
1). Next, it flows within
a narrow valley through
the Rożnowskie Foothills,
where it receives its largest
left tributary, the Łososina.
It cuts across Winnickie Figure 1. Location of the Czorsztyński Dammed Reservoir
Foothills and flows into the in the Dunajec River system.
Sandomierska Valley, where it joins its last big tributary, the Biała Tarnowska River. The
Dunajec, 247 km in length, drains water from a basin covering 6,804 km2 (in Poland,
4,852 km2). The average annual water flow in Nowy Targ is 14.3 m3.s-1, to increase to 84.3
m3.s-1 at the river’s outflow (Dynowska and Maciejowski 1991). In 1941, the valley of the
Dunajec River was crossed with a concrete water dam in Rożnowo, and in 1949 a smaller
dam was built in Czchów. Two reservoirs were created: a lager one called the Rożnowski
Reservoir, 1,600 ha in surface area, and a smaller one, an equalization reservoir, covering
346 ha. Quite recently, in 1997, on the border between Podhale and the Pieniny, two
139
other reservoirs were built: the Czorsztyński Reservoir (1,226 ha) and the Sromowiecki
Reservoir (88 ha) (Fig. 1). The water reservoirs Czorsztyn-Niedzica and Sromowce Wyżne
alongside the reservoirs in Rożnowo and Czchów are the main components of the water
economy over an area larger than Podhale. All these reservoirs constitute a system of
water resources management in the upper Vistula River and the water economy in its
basin. In 1996, a water power plant was started up in Niedzica, and a year later, in July
1997, the Czorsztyński Dammed Reservoir was officially opened. The reservoir was cre-
ated by raising the waters in the Dunajec River with a ground dam up to the height of
56 m. The length of the reservoir is 10.5 km, the surface area 1,226 ha and the capacity
is 231.9 million m3. The area of the drainage basin in the cross-section of the dam is
1,147 km2 and is characterized by quite a high altitude (415–2630 m above sea level) as
well as large annual precipitation, on average 1,072 mm in the whole basin. The average
annual water flow in the Dunajec River at the entry to the reservoir is 23.8 m3s-1. At the
water-raising site, there is a flow pumped-storage power plant of 92 MW.
The ichthyofauna of the Dunajec River before the reservoir was filled up with
water
Some intensive research on the ichthyofauna in the Dunaje River drainage basin ha been
going on since the 1950s, although the earliest reports date back to the late 19th century. The
first researcher who described the fish occurring in the Dunajec River was Nowicki (1883).
He distinguished the fish area of trout, which then covered the Dunajec River basin from
the river springs to Zakliczyn. In 1963-64, Kołder (1967) examined the ichthyofauna in the
Dunajec River from a drifting boat. In 1977-78, the ichthyofauna was investigated by Pas-
ternak and Skóra (1982), and before the Czorsztyński Reservoir was filled up with water, an
inventory of the ichthyofauna had been made by Augustyn and Bieniarz (1995).
In 1963, Kołder (1967) determined the presence of 12 fish species in the reservoir’s
bowl, among which the following were dominant in the abundance: nase Chondrosto-
ma nasus (30.59%), barbel Barbus barbus (26.03%) and spotted barbel Barbus carpathi-
cus (10.96%) (Kotlik et al. 2002) (Tab. 1). In terms of the biomass, the leading dominance
was created by barbel (44.58%) and nase (38.59%). In some later research, in 1977, nine
species were determined (Pasternak, Skóra 1982). In respect of the abundance, nase
continued to dominate (38.14%), but the second position of barbel had been taken over
by spotted barbel (23.73%) and bleak Alburnus alburnus (L.) (11.86%). As for the bio-
mass, nase was dominant (65.36%) (Fig. 2). In another study run in 1994 along a 10.5-
km-long section of the Dunajec within the future reservoir’s bowl, 17 fish species were
found (Augustyn and Bieniarz 1995). At that time, chub was dominant both in respect of
abundance (52.52%) and biomass (74.52%). Significant quantitative changes (Kruskal-
Wallis test X2emp = 27.56, P<0.0001) were accompanied by qualitative changes. When
the construction work on the reservoirs became more intensive (1981–1996), a large
decrease in the abundance of nase and barbel occurred. Their disappearance was related
to their foraging behavior. Foraging on periphyton algae and penetration of benthic
sediments makes these fish more vulnerable to contamination. Since 1983, there have
not been any water bodies with completely pure water found in the drainage basin of
the Dunajec River (Augustyn 2001). In an ichthyopathological study run in 1992, up
140
to 87% of nase and 41% of

barbel suffered from sep-
sis of cyprinid fish in the
form of highly extensive
and infectious erythroder-
matitis (Markiewicz 1992).
Meanwhile, the popula-
tion of chub, the most
omnivorous and eurytopic
fish, evidently grew larger.
The position of common
bream and roach increased
and became stable, which
was connected with their
transfer from gravel pits
created in the future reser-
Figure 2. Changes in the structure of the Dunajec River ichthyofauna
voir’s bowl (Augustyn and before filling the Czorsztyński Dammed Reservoir.
Bieniarz 1994).
Table 1. Composition and structure of the ichthyofauna in the Dunajec River within
the future Czorsztyński Dammed Reservoir in different years, acc. to: Kołder (1963),
Pasternak (1964, 1977), Skóra (1982), Augustyn (1995), Bieniarz (1995).
1963 1977 1995

Species Species % % %
(common name) (sciencific name) % % %
abun- abun- abun-
mass mass mass
dance dance dance
Brown trout Salmo trutta 4.11 2.06 3.39 3.19 4.36 3.12
Carpathian sculpin Cottus poecilopus 0.26 0.04
Stone loach Barbatula barbatula 1.79 0.07
Common minnow Phoxinus phoxinus 1.28 0.03
Rainbow trout Oncorhynchus mykiss 0.91 0.31
Chub Leuciscus cephalus 7.76 7.00 6.78 4.65 52.52 74.52
Spotted barbel Barbus carpathicus 10.96 2.12 23.73 6.43 14.01 3.52
Spirlin Alburnoides bipunctatus 3.65 0.14
Gudgeon Gobio gobio 3.39 0.16 1.37 0.12
Bullhead Cottus gobio 0.91 0.05
European grayling Thymallus thymallus 8.22 4.20 5.08 3.79 3.59 2.88
Huchen Hucho hucho 0.08 0.04
Barbel Barbus barbus 26.03 44.58 6.78 13.87 9.56 4.27
Dace Leuciscus leuciscus 1.37 0.23 0.34 0.19
Nase Chondrostoma nasus 30.59 38.67 38.14 65.36 1.02 1.75
Perch Perca fluviatilis 1.96 0.97
Roach Rutilus rutilus 0.46 0.17 0.85 0.34 4.01 1.21
Common bream Abramis brama 0.43 3.44
Bleak Alburnus alburnus 2.74 0.43 11.86 2.21 0.94 0.10
Pike Esox lucius 2.48 3.73
141
Fishing and angling management and exploitation of the Czorsztyński Reservoir

Pasternak and Skóra (1982) were the first to suggest some guidelines for the fish
resources management in the then constructed Czorsztyński Dammed Reservoir. For the
primary cyprinid species, they proposed vimba Vimba vimba carinata, and for the main
predator – rainbow trout Oncorhynchus mykiss. Other supported species should be – ac-
cording to these authors – roach Rutilus rutilus, bleak Alburnus alburnus, ide Leuciscus
idus and phytophagous nase. They also suggested that such species as common bream
Abramis brama and perch Perca fluviatilis should be eliminated.
In June 1989, a symposium was held in Niedzica named “The Dunajec River – yes-
terday – today – tomorrow”. The general principles for the management of the future
Czorsztyński Reservoir were then formulated by Kownacki and Starmach (1989). The
top priority was to maintain right proportions between particular fish species. The
main treatment suggested for maintenance of the proper structure of the ichthyofauna
should consist of fishery catches with dragged tools. Another conference called “The
current state and prospects of the ichthyofauna in the Dunajec River drainage basin”
was organized in Łopuszna by the Polish Angling Association, Region in Nowy Sącz.
A new concept for the reservoir’s fishing management was presented by Sych (1992),
who proposed lake trout Salmo trutta lacustris to act as a key predator. This concept,
enriched by adding another species, European whitefish Coregonus lavaretus (L.), was
approved of at the 8th meeting of the Working Group on Fisheries in Dammed Reservoirs,
affiliated at the Board of Directors of the Polish Angling Association in Warsaw (Anony-
mous 1994a). Further programme discussion continued during the 2nd meeting of the
Scientific Board on Fisheries Management in the Dunajec River drainage basin on 26
October 1994 (Anonymous 1994b), adding another species to the fishing and angling
plan, namely huchen Hucho hucho. As can be seen from the above, concepts for the
fishery management of the reservoir evolved from an idea of production intensification
(Kownacki and Starmach 1989) to an ecological way of managing the fish resources with
lake trout and huchen playing the role of large fish-eating predators which would be
able to stabilize the development of cyprinid species.
The gravel pits in the reservoir’s bowl remained a problem, as a specific structure
of the ichthyofauna was forming in those water pools (Augustyn and Bieniarz 1994).
The newly opened reservoirs were first colonized by pike, which nonetheless was un-
able to produce numerous populations. It was not only until a reservoir was colonized
by roach, which typically happened in the third year after the gravel exploitation had
terminated, that the development of the pike population could accelerate and peak in
the fifth or sixth after the reservoir had been filled up. The riverine species and the perch
did not show any significant changes in abundance. After ten years, the gravel pits were
dominated by common bream, which made up over 80% of the biomass. This succession
implies that there is a relationship between populations of pike and roach. The resources
of roach have a limiting effect on the abundance of pike. Similar mechanisms involved in
the formation of ichthyofauna have been described to take place in numerous dammed
reservoirs (Holčik 1966, Starmach 1986, Kubečka 1993, Řiha et al. 2009)
Because that fish from the gravel pits spread to the reservoir when it was being
filled up with water and put at risk the execution of the concept of fishery manage-
142
ment based on lake trout and huchen as key predators, in 1995 over 24 thousand
fish of the total weight of over 8 tons were caught from these pits using fishing nets
and electrocatches (Tab. 2). When the fishery management was re-directed so as to
prefer salmonid fish, the status of the waters in the Czorsztyński Dammed Reservoir
was changed from lowland to montane one. Pike Esox lucius (L.), which until then had
played a special role in the division of food resources, by regulating the number of prey
and thus maintaining the whole fish assemblage in a state of relative equilibrium, lost
any protection. It is estimated that since the ice cover thaw to the end of 1998, over
6 tons of pike was caught, mainly comprising spawners entering the Dunajec River in
search of spawning grounds.
Table 2. Composition and structure of fish captured from water-filled gravel pits before filling
the bowl of the Czorsztyński Dammed Reservoir.
Species n (indiv.) % (indiv.) mass (kg) % mass

Leuciscus cephalus 130 0.53 51 0.63
Tinca tinca 240 0.98 8 0.10
Carassius carassius 482 1.97 201 2.49
Chondrostoma nasus 23 0.09 18 0.22
Perca fluviatilis 1250 5.12 85 1.05
Rutilus rutilus 8630 35.36 1911 23.69
Abramis brama 8406 34.44 4461 55.29
Alburnus alburnus 2500 10.24 25 0.31
Esox lucius 2747 11.25 1308 16.21
Total: 24408 100 8068 100
Table 3. Fish stocking of the Czorsztyński Dammed Reservoir in 1996-2008.
Hucho hucho Salmo trutta lacustris Sander lucioperca Esox lucius
summer autumn
Year summer fingerling larvae smolt summer spawners
fry fry (kg)
fry (indiv.) (indiv.) (indiv.) (indiv.) fry (indiv.) (kg)
(indiv.)
1996 250000
1997 400000
1998 300000 13000 200000
1999 420000 70000 25000
2000 200000
2001 200000
2002 120000 132 3000
2003 45000 800
2004 63600 750
2005 60000 445
2006 45000 550
2007 36000 0
2008 105000 1000
Total 1890000 13000 270000 25000 354600 132 3000 3545
143
As early as in 1993, a batch of 5 thousand eyed-eggs of lake trout Salmo trutta lacus-
tris were brought from the fish hatchery in Wdzydze in order to make a maternal herd.
Thus, it was possible to initiate fish releases even before the reservoir was completely filled
up with water. Until 2002, the Czorsztyński Reservoir had been stocked with 1.89 million
individuals of larvae and 13 thousand fish fry of huchen as well as 270 thousand individu-
als of larvae and 13 thousand individuals of smolt of lake trout (Tab. 3). The lack of any
evident success of stock releases of salmonid fish caused the programme to discontinue.
The method of tagging fish with Carina type tags was adapted. In spring 2002, 1,648
individuals of tagged smolts of lake trout, 1,315 individuals of brook trout at the age cor-
responding to the smolt (2+) as well as 651 indiv. of huchen selects were released.
Until the end of 2003, only one return of brook trout was observed, when the fish
emigrated from the artificial lake 9 km upwards the Dunajec to Waksmund. The lack of
the expected results proved that the concept for future exploitation of the Czorsztyński
Reservoir had to be changed. It was therefore agreed that the assumed level of preda-
tory pressure necessary for the maintenance of the proper dynamic balance in the fish
assemblage would be provided by pike, pike-perch Sander lucioperca and, later, catfish
Silurus glanis. Until the end of 2008, the reservoir had been stocked with 654 thousand
of fry and 132 kg of spawners as well as 3 thousand individuals of summer fry and 3.5
tons of autumn fry of pike (Tab. 3).
Formation of the fish assemblage in the reservoir in the light of registered an-
gling catches
It is obvious that exploitation of fish, by definition selective, is responsible for chang-
es in assemblages of fish, but it also shares the responsibility for any reconstruction of
the structure of fish populations. In the spring of each year when the reservoir was be-
ing filled up with water (1993-1997), pike entered freshly flooded meadows, where they
found excellent conditions for natural reproduction. This migratory behavior was charac-
teristic for the whole pike population in the Czorsztyński Reservoir. The mass catches of
fish, mentioned earlier, which took place in 1997-1999 led to a collapse in the structure
of the ichthyofauna so that this predatory fish practically disappeared from the reser-
voir (Tab. 4). Since 2003, owing to fish releases of the stock material cultured in ponds,
the pike population has been partly replenished. However, in terms of the phenotype
and behaviour, it is a different type of pike. It does not migrate up the tributary riv-
ers for spawning, nor does it search for meadows, but chooses the rocky shores with
small boulders as an alternative spawning ground in the reservoir. Such behavior in the
natural reproduction has also been reported to occur in the Rimov Reservoir (Hladik and
Kubečka 2004). The size of captured pike individuals has also changed. In 1999-2000
small fish up to 65 cm in length dominated. At present, the catches are dominated by
larger pike specimens, between 65-85 cm long (Fig. 3).
Now, next to pike-perch, pike is the basic component of angling catches of predatory
fish. The first catches of pike-perch were recorded in 2004, and kept increasing nearly
geometrically in the following four years (Tab. 4). Likewise, there has been a progress,
albeit a small one, in the catches of perch. Until present, the Czorsztyński Dammed Res-
ervoir has not been observed to be the site of a highly dynamic and unstable dominance
144
of perch. In this respect, this reservoir is different from other artificial lakes of this type
(Starmach 1986, Řiha et al. 2009).
The study conducted by Starmach and Jelonek (2003) showed that the shortening
of the trophic chain had led to a specific population explosion of roach, previously re-
stricted by the predatory pressure exerted by pike. In 2000-2001, roach made up 58%
of the total amount and 34% of the biomass of the ichthyofauna in the Czorsztyński
Reservoir. Angling catches
of roach exceeded 4.5 tons
in 2000, and it was not un-
til more angling pressure
was directed onto com-
mon bream that the game
catches of roach declined
(Tab. 4). In later years,
catches of roach have
Figure 3. Size distribution of pikes caught by anglers
been increasing and now
in the Czorsztyński Dammed Reservoir.
are over 6.5 tons a year.
Table 4. Angling catches (kg) in the Czorsztyński Dammed Reservoir in 1999-2008 (Augustyn 2010).
Species / Year 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008
Salmo trutta fario 27.4 3.3 6.0 2.0 1.1 2.1 1.4 5.3 62.6
Salmo trutta lacustris 41.6 4.8 63.7 18.2 17.1
Oncorhynchus mykiss 1.2 44.6
Thymallus thymallus 19.4
Hucho hucho 9.0 9.4 21.5 13.6 19.6 28.2
Barbus barbus 2.3
Chondrostoma nasus 53.3 2.5 25.2
73.6
Leuciscus cephalus 181.5 449.8 300.6 46.4 71.5 158.9 159.2 128.4 220.4
581.7
Leuciscus leuciscus 2.4 0.5 0.5
0.8
Leuciscus idus 2.2
Esox lucius 3270.7 1916.8 222.8 60.6 233.5 172.6 321.4 658.7 563.7 829.8
Sander lucioperca 4.8 20.3 340.8 376.0 1538.9
Perca fluviatilis 587.3 1808.8 1134.4 1808.1 735.2 2317.6 1021.2 4007.6 2962.6 5318.7
Anquilla anquilla 1.8 2.3
Aspius aspius 1.7
Abramis brama 106.3 1902.8 2174.6 10360.1 8948.9 6727.6 7118.8 7868.9 8765.4 1677.6
Carasius carasius 0.4 8.0 1.8 15.8 0.4 1.3
Tinca tinca 24.3 15.5 8.3 46.8 5.1 1.8 1.7 3.2
Rutilus rutilus 2017.3 4529.7 3734.7 2806.5 2474.5 5587.4 6229.4 5229.6 5645.3 6526.8
Alburnus alburnus 14.2 15.7 26.7 35.7 16.3 4.9 16.9 39.1 12.9
Cyprinus carpio 1.8 26.5 11.4 14.0 12.1
Gymnocephalus cernuus 0.1 0.4 0.1
Blicca bjoerkna 29.1 6.8
TOTAL 6190.5 10705.5 7626.5 15120.5 12571.1 15196.3 14928.8 18291.2 18638.8 16705.6
In the initial stage of the formation of the ichthyofauna in the Czorsztyński Reser-
voir, the most dynamic species was common bream, whose older specimens originating
from spawning events in the gravel pits dominated game catches. However, since 2003
145
catches of this species have been declining and the captured specimens tend to be
smaller (Fig. 4). Why does it happen? In many reservoirs, populations of common bream
grow dynamically until acquiring complete dominance (Starmach 1986, Wiśniewolski
2002, Řiha et al. 2009). Al-
though common bream is
highly tolerant to environ-
mental conditions and is
an omnivore, in contrast
to roach or perch, it is not
a fish with a short life-cycle.
The factor that hinders the
development of its popu-
lation in the Czorsztyński Figure 4. Size structure of bream, caught
Dammed Reservoir is the in the Czorsztyński Dammed Reservoir in 1999–2008.
complete absence of macrophytes in the littoral zone. A similar situation occurred in the
Rożnowski Reservoir, where twenty years after the lake had been filled up with water,
common bream did not play any important role in commercial catches (Augustyn 2001).
It was not until plants had grown in backwater areas in the reservoir that common
bream could achieve reproduction success to the point where completely colonized the
reservoir in the 1980s (Bieniarz et al. 1990).
Another factor that limits the reproduction potential of common bream in the
Czorsztyński Reservoir are fluctuations in the level of the water table, especially in spring.
Common bream in the Czorsztyński Reservoir starts spawning rather late, from the end
of May to mid-June. Changes in the water table level during that time affect the acces-
sibility of spawning grounds and therefore the reproduction success.
Summary
Formation of ichthyofauna in dammed reservoirs is usually divided into several phas-
es (Starmach 1986, Wiśniewolski 2002, Řiha et al. 2009). The first phase consists of the
dominance of riverine species, which have been flooded by the raised waters of the
dammed reservoir. This phase ends after a few years when river species leave the res-
ervoir. However, they sometimes form stable populations, which dwell in the reservoir
but enter rivers for spawning. Examples are the vimba and nase in the Orawski Reservoir
(Holčik 1966), vimba and nase in the Rożnowski Reservoir (Bontemps 1968, Augustyn
2001) or chub in the Klimówka Reservoir (Augustyn et al. 2003). The same situation
occurs in the Czorsztyński Reservoir, where at present catches of nase and chub are on
the increase. In the first stage after a reservoir is filled up with water, its trophic state
increases because the flooded soils are leached off from large amounts of mineral nutri-
ents (Starmach 1986), which limits the abundance of rheophilic species. Afterwards, the
fertility of the reservoir decreases and it can be colonized by riverine species less tolerant
to the environmental pollution.
Any further fish status in a dammed reservoir depends to a large extent on the re-
lationship between the drainage basin and the reservoir itself. Anthropogenic factors,
mainly area and point pollution, play the most important role here. The fish assemblages
146
that are being formed, colonize a new environment and are subjected to accelerated
succession. In the majority of dammed reservoirs, during the second phase, a popula-
tion explosion of pike appears. In the Czorsztyński Reservoir, this stage was consider-
ably shortened because any passive conservation of this species had been abandoned.
This, however, did not lead to a rapid growth in the number of common bream in the
reservoir. Fish represent the highest trophic level in aquatic ecosystems, which makes
them responsive to a variety of environmental disturbances. It is not, however, an im-
mediate response but a delayed one relative to the lower trophic levels. The Czorsztyński
Reservoir had been filled up with water before the water and sewage management in
the basin was completely put into order. The changes in the angling catches of species
indicating eutrophication of lakes (common bream), discussed in this paper, prove that
the process of an increase in the trophic state of the reservoir has already been halted.
Results of some research confirm this conclusion. In 2004, following some changes in
the land use in Podhale, nitrogen loads from large-area pollution fell down 11-fold over
fifteen years (Kopacz and Twardy 2006). The evident improvement in the quality of wa-
ters in the upper Dunajec River has been aided by a crisis in the manufacturing of fur
clothes (illegal fur processing) (Szalińska and Dominik 2006). Is it therefore possible to
return to the concept of stock releases with huchen and lake trout? Certainly not! The
formation of the ichthyofauna in the Czorsztyński Reservoir runs according to the gener-
ally known patterns. In this case, the imperfection of our scientific knowledge has been
painfully verified by nature.
It is generally believed that recreational angling creates many risks, mainly exces-
sive dilution of populations of attractive game species of fish, or the elimination of top
predators from the environment (Arlinghaus and Cooke 2005). This opinion, generally
held true in the world, does not find confirmation in Poland, especially among anglers
catching fish in artificial lakes, who prefer roach and common bream, fish species which
do not require high angling skills. Most anglers treat fishing as a way of obtaining food
(Wrona and Guziur 2006). In this context, angling catches serve a typical function of
a regulator of ichthyofauna assemblages (Wołos 2000, Draszkiewicz-Mioduszewska and
Wołos 2007, Wołos 2008).
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149
Structure of ichthyofauna in
different fishery types of lakes
used for angling by the Polish
Angling Association, the Region
of Szczecin
Adam Tański1, Przemysław Czerniejewski2,
Sławomir Keszka3, Rafał Pender4
1
Department of Fish Anatomy and Embryology, Faculty of Food
Sciences and Fisheries, e-mail: atanski@zut.edu.pl
2
Department of Sustaiable Development of Fishing Industry
Management, Faculty of Food Sciences and Fisheries,
e-mail: pczerniejewski@zut.edu.pl
3
Fish Taxonomy Unit, Faculty of Food Sciences and Fisheries,
e-mail: skeszka@zut.edu.pl
West Pomeranian University of Technology in Szczecin
4
Polish Angling Association, Region in Szczecin,
e-mail: rafal@pzwszczecin.com
Abstract
The Polish Angling Association, the Region in Szczecin, manages over 2,649 ha of lakes
for fishing and angling. The annual fish stocking and angling exploitation cause changes in
the structure of ichthyofauna. Based on the control catches performed in 13 lakes, which be-
long to the following fishery types: tench-pike, bream, pikeperch and vendace, an analysis of
the structure of ichthyofauna in lakes used for angling has been made. The results indicate
that the fish catches are dominated by cyprinid fish species (60-80% of captured fish), and
that particular fishery types of lakes differ in respect of the share of predatory fish.
Introduction
The Polish Angling Association Region of Szczecin is one of the 47 regions of the PAA
in Poland. It has 14,080 members (in 2008). The primary objectives of the association are
expressed in the organization’s statute. The overriding aim is conservation of the natural
environment by rational management of fishing and angling activities, i.e. fish stocking
151
of water bodies, sustainable fish catches, protection from pollution and poaching. The
PAA Region in Szczecin manages 2,649 ha of lakes, 1,291 ha of rivers, 283 ha dam reser-
voirs and 28 ha of canals. These water bodies are stocked every year with different fish
species, e.g. in 2008 the fish stock releases cost 756,000 Polish zloty (Tański and Pender
2008). Parallel to this and apart from angling catches, selective, control and spawner
catches are carried out (Tański and Pender 2009).
Recently, angling has been one of the most rapidly developing forms of exploitation
of open waters. On the one hand, it is an excellent hobby for many people, being one
of the most popular forms of relaxation and recreation. On the other hand, it is a seri-
ous branch in the fisheries, which has a considerable influence on the structure of the
ichthyofauna composition. The current number of anglers largely exceeds the number of
the PAA members and is estimated to be as high as 2 million people (Wołos 1994). Ow-
ing to its mass character and intensive fish catches, angling may contribute to excessive
scarcity of some fish species exploited in water bodies. When they are not stocked, the
situation can lead to selective limitation of some species or even size (age) classes of fish,
while other fish are not caught in appropriate amounts. In an extreme case, the whole
fish resources can be overfished. This event may be a cause of undesirable modifications
in fish resources (Leopold and Bnińska 1987). Thus, the water bodies managed by the
PAA Region of Szczecin underwent a study, the aim of which was to determine the struc-
ture of ichthyofauna in lakes belonging to different fish lake types.

Table 1. Specification of lakes exploited by the PAA Region of Szczecin in which control catches were run.
Name of a water Fishery Area Depth [m]

Date of catches
body type [ha] max/average
Bielin bream 32.7 12.0/5.2 18.11.2009
Chociwel pikeperch 59.2 6.1/2.6 11.09.2009
Głębokie k. Radowa bream 41.7 11.5/4.3 28.11.2009
Klasztorne tench-pike 16.3 3.1/1.5 7.07.2009
Kołomąć tench-pike 39.1 4.6/2.6 17.07.2009
Morzycko vendace 342.7 62.0/16.5 18.11.2009
Nowogard bream 98.3 10.9/5.1 5.06.2009
Piaski tench-pike 28.6 6.8/2.1 30.06.2009
Przybiernów bream 88.9 5.3/2.76 21.05.2009
Stara Dobrzyca bream 29.8 8.2/4.5 7.05.2009
Trzcińsko pikeperch 43.6 5.6/3.1 3.07.2009
Trzygłów I pikeperch 17.4 11.9/5.5 30.07.2009
Trzygłów II tench-pike 15.3 7.3/3.9 21.07.2009
The catches were performed in 2009-2010, in 13 lakes (Tab. 1) managed by the PAA
Region of Szczecin. The catches were done using 10 gillnets, each 50 m long. The mesh
sizes in the gillnets were: 10mm, 22mm, 35mm, 55mm and 70mm. Apart from net
152
Structure of ichthyofauna in different fishery types of lakes used for angling…
fishing, fish were also caught in the littoral zone from a boat, by electrocatches, using
a diesel oil energy generator, catching fish along 150-meter-long sections. Catches were
conducted from 6.00 p.m. to 6.00 a.m.. Afterwards, the species identity of captured fish
was determined along with their number and total weight within each species. Moreo-
ver, the lakes where the fish were caught were divided into fishery types, and the species
structure of the ichthyofauna was established.
Results
Among the captured fish, 14 species were identified, predominantly cyprinid ones (8
species). The data found in fishing valuation surveys enabled us to classify the lakes into
fish lake types, between which considerable disproportions in the number of fish species
inhabiting these types of lakes were found.
During the catches in the pikeperch type of lakes (Trzygłów I, Trzcińsko, Chociwel),
239 fish were captured (Tab. 2). The largest biomass in the catches was attributed to
bream (45.48%), and pikeperch was the second species in respect of the total biomass
in these lakes (Fig. 1).
Table 2. Species and qualitative structure of fish caught during control catches in particular lakes
of the PAA Region of Szczecin (the data in the table regarding particular species comprise numbers
of captured fish and their total weight expressed in grams [indiv./g])
Individuals [indiv./g]
pikeperch
prussian
vendace
crucian
In total
catfish
Name of lake
bream
roach
perch
tench
rudd
carp
carp
carp
carp
pike
eel
23/ 31/ 58/

Bielin 2/239 - - 2/2779 - - - - - - - -
30287 17456 50761
33/ 18/ 21/ 80/
Chociwel - - - 2/1278 1/767 2/7752 - - 2/5773 1/397 -
59562 56021 22072 153622
17/ 25/ 48/
Głębokie - - 3/425 2/2463 - - - - - 1/2120 - -
1533 5210 11752
9/ 49/ 83/
Klasztorne 5/1388 - 4/444 5/1976 - - - - 3/2747 - 8/2442 -
6209 24799 40005
47/ 60/
Kołomąć 3/511 1/372 1/57 3/554 2/7921 - - 2/404 - - - 1/621 -
40611 51051
15/ 10/ 33/
Morzycko 2/180 - - 1/500 1/1762 1/1331 - 2/2088 - 1/1345 - -
21164 901 29271
45/ 61/
Nowogard - - - 4/499 5/8349 - - - - 5/3800 1/2458 2/691 -
51620 64959
23/ 36/
Piaski - 1/105 - 4/644 3/5956 - - - 3/4447 - 2/796 -
12646 24594
80/ 43/ 21/ 4/ 8/ 191/
Przybiernów 14/960 9/409 7/4564 - - - 2/7059 3/866 -
2420 1247 6441 30300 2904 57173
28/ 11/ 61/
Stara Dobrzyca - - 7/2193 2/1690 - - - - - - - -
3310 1368 8562
7/ 21/ 31/ 8/ 16/ 95/
Trzcińsko 1/302 3/1345 4/3465 - - - 4/2922 - -
1484 2938 24346 5776 12389 54967
9/ 39/ 64/
Trzygłów I - 8/1747 2/1963 4/4130 - - - 1/765 - 1/686 -
1005 28683 38979
39/ 11/ 4/ 2/ 10/ 73/
Trzygłów II 3/581 - 4/660 - - - - - -
42743 2323 7311 14193 7148 74959
153
In bream lakes (Now-

ogard, Przybiernów, Bielin,
Głębokie near Radowo,
Stara Dobrzyca), 419 fish
were caught, representing
10 species and weighing
193,207 g in total (Tab. 2).
In these lakes, the highest
weight percentage was
that of bream (48.95%)
and predatory fish such Figure 1. Structure of the biomass of ichthyofauna in pikeperch lakes
as catfish (15.49%), perch (Trzygłów I, Trzcińsko, Chociwel).
(10.28%) and pike (10.15%)
(Fig. 2).
Lakes classified as the
tench-pike type (Klasz-
torne, Piaski, Trzygłów II,
Kołomąć) were character-
ized by a larger number of
captured species than the
former two lake types. In
total, 12 fish species were
obtained, among which the Figure 2. Structure of the biomass of ichthyofauna in bream lakes (Now-
ogard, Przybiernów, Bielin, Głębokie near Radowo, Stara Dobrzyca).
highest biomass was made

up by bream (53.62%) (Fig.
3). Moreover, 13% of the
biomass of the fish caught
in these lakes consisted of
prussian carp.
The vendace lakes were
represented by one lake,
Morzycko Lake, in which
Figure 3. Structure of the biomass of ichthyofauna in tench-pike lakes
(Klasztorne, Piaski, Trzygłów II, Kołomąć). 33 fish belonging to 8
species were caught (Tab.
2). The dominant species
was bream (72.30% of the
biomass of captured fish)
and then eel (7.13%), pike
(6.02%), tench (4.59%) and
pikeperch (4.55%). Ven-
dace made up 3.08% of
the mass of captured fish
Figure 4. Structure of the biomass of ichthyofauna in vendace lakes
(Morzycko Lake). (Fig. 4).
154
Structure of ichthyofauna in different fishery types of lakes used for angling…
Discussion
West Pomerania is an important region in respect of fishing, owing to a large number
of lakes and rivers, used for organized fisheries and angling. According to Choiński
(2006), the whole region has 3,385 lakes, which in total cover around 104,219 ha. This
is quite a large number of lakes, considering that Poland has 7,085 lakes in total, as
listed in the Catalogue of Polish Lakes by Choiński (2006). Filipiak and Raczyński (2000)
report that the Province of West Pomerania (województwo zachodniopomorskie) alone
has 1,575 lakes over 1 ha in area. According to a study performed by Czerniejewski et
al. (2003) on 1,413 lakes, the dominant lakes in this region are crucian carp (513) and
tench-pike lakes (508), of the total area of 2,038.6 ha and 11,204.6 ha, respectively.
The pikeperch type of lakes comprises 249 lakes, covering in total 30,360.8 ha, with the
average acreage of 121.9 ha. Among the lakes in the Province of West Pomerania, there
are 59 vendace and bream lakes, covering in total 18,157.5 ha and 7,505 ha, respec-
tively. The vendace-bream subtype of lakes is represented by 18 lakes of the total area
of 2,773.2 ha, whereas the bream-pikeperch type encompasses 7 lakes of the total area
of 332.5 ha.
This classification, although based on morphometric and environmental parameters
as well as leading fish species in the ichthyofauna, gives only a general understanding
of possible productivity of lakes and a potential share of individual fish species in fish
catches, as a result of appropriate fish management treatments. The above classifica-
tion, alongside theoretical fish catching potential, is presented in Table 3.
Table 3. Theoretical fish yields for different fishery types of lakes (Szczerbowski 1993).
Fishery types of lakes
Species vendace bream pikeperch tench-pike crucian carp
estimated fish yield (kg/ha)

vendace 5.5 - - - -
whitefish 0.5 - - - -
tench 5.0 5.0 5.0 8.0 2.0
pike 1.0 3.0 1.0 4.0 3.0
eel 1.0 1.0 2.0 2.0 1.0
bream 3.0 6.0 6.0 - -
pikeperch - 0.5 4.0 - -
crucian carp - - - 2.0 6.0
perch 2.0 3.0 3.0 4.0 0.5
roach 3.0 3.0 3.0 4.0 2.0
white bream 3.0 4.0 4.0 4.0 1.0
others* 10.0 8.5 10.0 12.0 4.5
In total 34.0 34.0 38.0 40.0 20.0
*/ catfish, blue bream, chub, ide and fish species with no-economical value
155
In the lakes used for angling, there is no regular net fishing, which is why it is impos-
sible to determine the fishing yield. However, owing to the fact that angling catches are
usually selective ones, focusing on fish species that are more valuable to anglers, the
so-called control net fishing catches should be carried out periodically. Such catches
will show the current state of fish resources in a water body. Our analysis of the control
catches performed in 13 lakes has revealed evident dominance of cyprinid fish, whose
share ranged from 60 to 80% depending on a fishery type of lakes. Large disproportions
have been recorded in the share of predatory species between particular fishery types of
lakes. The highest share of pikeperch was noticed in pikeperch lakes (26.63%), while in
the other analyzed lake types, it did not exceed 5%. The percentage of pike varied from
2.5% (in pikeperch lakes) to 10.15% (in bream lakes). Another interesting finding is the
high share of catfish in bream lakes (15.49%). In the other types of lakes this fish was
unobserved except pikeperch lakes (3.13%).
References
Choiński A. 2006. Katalog jezior Polski. Wydawnictwo Naukowe UAM. Poznań: 600 pp.
Czerniejewski P., Raczyński M., Filipiak J. 2003. Rybackie typy jezior w województwie zachodniopo-
morskim. Materiały Ogólnopolskiej konferencji naukowej „Stan badań naukowych, jakości wód i prak-
tyki rybackiej przed wejściem Polski do Unii Europejskiej”. Międzyzdroje 26-28 czerwca 2003: 100 pp.
Filipiak J., Raczyński M. 2000. Jeziora zachodniopomorskie. Wyd. AR Szczecin.
Leopold M., Bnińska M. 1987. Ocena presji połowów wędkarskich na pogłowie poszczególnych
gatunków ryb w wodach Polski – konsekwencje gospodarcze. Rocz. Nauk. Rol. 2: 43–69.
Szczerbowski J. A. 1993. 1993. Rybactwo śródlądowe. Wyd. IRS Olsztyn: 570 pp.
Tański A., Pender R.. 2008. Charakterystyka gospodarki wędkarskiej prowadzonej przez Okręg
Polskiego Związku Wędkarskiego w Szczecinie. Rybackie perspektywy pobrzeża południowego
Bałtyku, tom II, Wyd. FOKA, Szczecin: 73-82.
Tański A., Pender R. 2009. Gospodarka rybami łososiowatymi prowadzona przez Okręg Polsk-
iego Związku Wędkarskiego w Szczecinie w latach 1999–2009. In: Rozród, podchów, profilaktyka
ryb łososiowatych i innych gatunków [Eds. Zakęś Z. Demska-Zakęś K., Wolnicki J ]. Wyd. IRŚ, Olsz-
tyn: 243–250.
Wołos A. 1994. „Put and take” – nowy model łowiska wędkarskiego. In: Aktualne problemy
rybactwa jeziorowego. [Ed. Wołos A.], Wyd. IRS, Olsztyn: 133-139.
156
Biomanipulation as a method of
creating habitats for endangered
fish species: a case study of the
introduction of whitefish into
Lake Starodworskie
Bogdan Wziątek1, Renata Tandyrak2,
Magdalena Bowszys3
1
Department of Fish Biology and Breeding, University of Warmia
and Mazury in Olsztyn, ul. Oczapowskiego 5, 10-957 Olsztyn,
e-mail: b.wziatek@uwm.edu.pl
2
Department of Environmental Protection Engineering, University
of Warmia and Mazury in Olsztyn, ul. Prawocheńskiego 1, 10-957
Olsztyn, e-mail; renata.tandyrak@uwm.edu.pl
3
Department of Applied Ecology, University of Warmia
and Mazury in Olsztyn, ul. Oczapowskiego 5, 10-957 Olsztyn,
e-mail: mbowszys@uwm.edu.pl
Abstract
Biomanipulation is one of the methods applied to restoration of water bodies. Until
now, it has been used mainly to improve the quality of water in lakes or rivers of less
economic importance (water intakes, bathing beaches, etc.). Biomanipulation has not
yet been considered as a method to protect more sensitive fish species.
This study has covered Lake Starodworskie (surface area 6 ha, maximum depth 23.3
m), situated on the campus of the University of Warmia and Mazury in Olsztyn. With
respect to its limnological characteristics, the lake is meromictic. Until 2005, during the
summer stagnation, the oxygenated layer of water did not reach lower than 3 meters
and the Secchi disc transparency varied from 30 to 50 cm. Whitefish had never been
a component of this lake’s ichthyofauna. In 2004-2006, the lake was regularly stocked
with pike-perch and asp. Besides, catfish (twice) and eel (one) have been released to the
lake. In autumn 2005, 210 individuals of whitefish fry, weighing on average 25 g, were
released. In 2006 and 2007, control catches of this species were performed. Six and ten
specimens were captured, respectively. The growth rate was tested. By comparing with
157
the reference data, the growth rate of whitefish in Starodworskie Lake should be con-
sidered as moderately fast.
Introduction
European whitefish (Coregonus lavaretus L.) is a declining species in Poland. The
main reason are transformations of the environment, stimulated by the broadly under-
stood anthropopressure and consequent acceleration of lake eutrophication processes.
As a result, the number of lakes inhabited by whitefish has decreased greatly over the
last 30 year (Szczerbowski 1993, Szczerbowski 2000). With this undesirable tendency, it
is possible that the species may soon disappear from the Polish ichthyofauna despite the
fact that its artificial breeding and reproduction methods have been developed.
Biomanipulation has been a supporting technique applied in lake restoration for
over twenty years now (Shapiro et al. 1975, Gulatti et al. 19990, Kasprzak et al. 2000).
Biomanipulation relies on reduction of zooplanktivorous fish, especially their young rep-
resentatives, which are not commercially fished and therefore more difficult to eliminate
from the ecosystem (Shapiro 1990). However, there are no reports on biomanipulation
as a method of active conservation of threatened fish species.
The purpose of this study was to test whether a biomanipulation treatment, leading
to enhanced water transparency, would enable us to successfully introduce whitefish,
thus creating an artificial habitat for this endangered fish species.

Starodworskie Lake is located within the administrative area of the town of Olsztyn.
The water table surface area is 6.0 ha, the maximum depth is 23.3 m, the average depth
is 9.0 m and the length of the shoreline is 900 m (Tandyrak et al. 2009).
Among the zooplanktivorous fish inhabiting the lake, the following species dominat-
ed until 2004: sun bleak (Leucaspius delineatus), roach (Rutilus rutilus) and small perch
(Perca fluviatilis), whereas the predatory fish were represented by pike (Esox lucius) and
larger specimens of perch.
The biomanipulation treatment consisted in introducing predatory species not native
to the lake’s ichthyofauna, such as pike-perch (Sander lucioperca), asp (Aspius aspius),
catfish (Silurus glanis) and eel (Anguilla anguilla) (Tab. 1). Reared up fish fry was used for
stocking the lake: summer and autumn fry of pike-perch, summer fry of asp, autumn fry
of catfish, and elvers of eel.
Table 1. Predatory fish fry doses stocked to Starodworskie Lake in the biomanipulation experiment.
Species Dose (indiv./ha) Year

pike-perch 330 2004-2006
asp 120 2004-2006
catfish 60 2005
eel 30 2004
During the experiment, the Secchi disc transparency of water was determined eight
times a year. Oxygen concentration in lake water was measured four times each year
using the Winckler’ s method.
158
Biomanipulation as a method of creating habitats for endangered fish species…
The abundance of zooplanktivorous fish in the lake was assessed with the CPUE
(catch per unit effort, Berg 1998) method. Control catches were run using 4 gillnets with
of the meshes 10 to 16 mm in size and 10 m long. The gillnets were raised every twelve
hours. All the captured fish were counted and weighed. The catches were repeated in
the three-days intervals. Based on the information thus gathered, the CPUE value for
a given year was calculated.
Whitefish was stocked into the lake in December 2005, using 215 individuals of
autumn fry, each of them weighting 21.5 g on average. Control catches of whitefish
were performed twice: in 2006 and 2007. Three sets of gillnets of the mesh size 35, 40
and 45 mm (2006) and 40, 45 and 50 mm (2007) were used. The catches ran for three
days. The captured fish were measured and weighed. Their scales were sampled to as-
sess the growth rate. All the manipulations, whenever possible, were performed on live
fish, which were afterwards released back. In total, 16 fish were caught: 6 in 2006 and
10 in 2007.
In order to standardize the data for statistical processing, the term of average ef-
ficiency was introduced. For this purpose, the results of the Secchi disc transparency
measurements made on particular days were averaged for a whole year and then cor-
related with the average results of the CPUE results.
The growth rate of whitefish was assessed on the basis of the readings from scales
and direct measurements of the fish. A model of the growth rate was drawn according
to a modified power function method (Szypuła 2000).
Results
Fish density
In 2004 (when the experiment was set up), the CPUE value was 269.5 and only one
fish species was captured, i.e. roach. In 2005, a considerable decrease in the number of
fish occuring in the lake was noticed: the CPUE value dropped to 147.5 fish/catch, but
in that year perch was caught apart from roach. In 2006, CPUE increased to 187.5 and
results revealed that perch began to dominate in the lake.
The decreasing value of CPUE was accompanied by an improvement of the transpar-
ency of water. In 2004, the average water transparency was 0.68 m, than rose to 2.1
m in 2005 and fell down again to 1.86 m in 2006. The values of the average water
transparency were strongly correlated with the abundance of fish (Spearman’s r = -1,
p<0.05). The highest value of 2.1 m was recorded in 2005, when the CPUE value was
the lowest (Fig. 1).
The growth rate of whitefish
In 2006, 6 specimens of whitefish were caught. Their average weight was 235.5 g
and the average length was 25.9 cm (Fig. 2). Among the ten whitefish captured in 2007,
males with visible spawn rash (6 individuals) predominated. Two of the fish were females
but the gender of two other ones could not be determined intra vitam. The average
weight of the fish was 517.2 g and the length was 31.9 cm (Fig. 3).
Considering the average lengths of the fish (l.c.) in the second (25.9 cm) and third
(31.9 cm) year of their life, the growth rate of whitefish in Starodworskie Lake should
159
be classified as moderately
fast (Fig. 4).
Discussion
Compared to other
Polish populations, the
growth rate of the white-
fish in Starodworskie Lake
should be regarded as
moderately fast when con-
sidering the body weight
and fast in respect of body
length (Szczerbowski 1981, Figure 1. Dependence between number of fish in the lake’s water and
2000). average Secchi disc transparency in Starodworskie Lake in 2004–2006.
Martyniak et al. (1999)

found out that the popu-
lation of whitefish inhab-
iting the Czarne Lake in
the Drawieński National
Park, which is also consid-
ered meromictic , similarly
to Starodworskie Lake,
was characterized by slow
growth in the length, even Figure 2. Two-year-old whitefish caught in Starodworskie Lake
though the fish did not af- in November 2006.
flicted with the competi-
tion of other fish species.
The growth rate deter-
mined for the population
from Starodworskie Lake
was comparable to that of
the population introduced
to Santa Croce Lake in Italy
(Loro and Mamcarz 2002).
Studies completed in Figure 3. Three-year-old whitefish caught in Starodworskie Lake
Wigierski National Park have in December 2007.
demonstrated that environ-
mental conditions, and especially improved water quality manifested by a rapid increase
in transparency and, consequently, better oxygen conditions in the meta- and hypolim-
nion, are very important factors which affect the growth rate of whitefish (Wziątek et
al. 2004). The steady growth rate of whitefish in Starodworskie Lake, as observed during
our experiment, suggests that regular predatory fish stocking stabilize the habitat condi-
tions, despite the angling and poaching pressure (own observations), so that no external
interference (regulation catches, etc.) is needed. That creates favorable conditions for
160
Biomanipulation as a method of creating habitats for endangered fish species…
whitefish development in
that lake despite the fact
that it had never inhabited
that lake before.
Until the present day, bi-
omanipulation treatments
have been conducted in
shallow lakes, lacking in
thermal stratification (Gu-
latti et al. 1990, Kasprzak et
al. 2002). They have rarely
been conducted successful- Figure 4. The growth of length (Sl) and weight of whitefish
in Starodworskie Lake.
ly in deeper water bodies,
where thermal stratification is evident (Lathrop et al. 2002). The occurrence of an anaero-
bic water layer above the bottom caused the transformation of phosphorus compounds
into dissolved forms, which again enrich the topmost water layers during the water circu-
lation period. Thus, the abundance and biomass of zooplanktivorous fish do not have any
significant effect on the quality water in such lakes (Lathrop et al. 2002).
With respect to Starodworskie Lake, the success of the experiment was most prob-
ably attributable to a small surface of the littoral zone, and thus the lack of sites where
young generations of zooplanktivorous fish could find a refuge, which exposed them to
stronger predatory pressure.
Another important factor was the meromictic type of the lake (Tandyrak and
Gawrońska 2009). Zooplanktivorous fish most probably affected the rate of phosphorus
cycling in the epilimnion (Shapiro et al. 1975). The decreased number of fish inhibited(!)
the timeover of phosphorus cycling in the epilimnion and the limnological type of the
lake, particularly the thermocline lying very shallow (3 m deep), prevented phosphorus
supply from deeper layers of the lake. (Tandyrak and Gawrońska 2009). This conclusion
is supported by observations on the range of oxygen limits in the lake. This parameter
reached its highest values at the end of summer stagnation (September), in contrast to
other lakes. It can indicate that as the summer stagnation continued in time, the nutri-
ents were being depleted in the epilimnion and the primary production was transferred
to deeper layers of the metalimnion.
These preliminary results seem to suggest that biomanipulation could be an effective
tool in protection of the threatened fish species of the genus Coergonus, especially in
lakes similar in character to Starodworskie Lake.
References
Berg S. 1998. Diet of rainbow trout (Oncorhynchus mykiss), stocked as a biomanipalution tool
in a eutrophic brackish lake. In: Stocking and introduction of fish. [Ed. Cowx, I.G.] Fishing. New
Books, Blackwell Science Ltd: 88-98.
Gulatti, R.D., Lammens, E.H.R.R., Meijer M.L., van Donk, E. 1990. Biomanipulation, Tool for
Water Management. Kluwer Academic Publishers, Dordrecht.
Kasprzak, P., Benndorf J., Mehner, T., Koschel, R. 2002. Biomanipulation of lake ecosystem: an
introdauction. Freshwater Biology 47: 2277-2281.
161
Lathrop R.C., Johnson B.M., Johnson T.B., Vogelsang M.T., Carpenter S.R. Hrabik T.R., Kitchell
J.F., Magnuson L.G., Rudstam L.G., Steward R.S. 2002. Stocking piscivores to improve fishing and
water clarity: a synthesis of the Lake Mendota biomanipulation project. Freshwater Biology 47:
2410–2424.
Loro R., Mamcarz A. 2002. Coexistence of introduced Perca fluviatilis and Coregonus lavaretus
in Santa Croce Lake (Italy), and proposition for fishery management. In: Management and ecology
of lake and reservoir fisheries. [Ed. Cowx, I.G.], Fishing New Books, Blackwell Science Ltd: 161-169.
Martyniak A., Terlecki J., Kozłowski J., Poczyczyński P., Wziątek B. 1999. Plan ochrony ichtio-
fauny, płazów, gadów, ptaków i ssaków związanych ze środowiskiem wodnym. Manuscript, the
Drawieński National Park.
Shapiro J. 1990. Biomanipulation in the next phase: making in stable. Hydrobiologia 200/201: 13-27.
Shapiro, J., Lamarra, V., Lynch, M. 1975. Biomanipulation: an ecosystem approach to lake resto-
ration. In: Water quality management trough biological control. [Eds. Brezonik P.L., Fox, J.L.], Report
no ENV-07-75-1 University of Florida: 85-96.
Szczerbowski J.A. 1981. Criteria of fish growth. Rocz. Nauk. Rol. 99: 123-136.
Szczerbowski J.A. 1993. Rybactwo śródlądowe. Wyd. IRŚ Olsztyn.
Szczerbowski J.A. 2000. Sieja. In: Ryby słodkowodne Polski [Ed. Brylińska M], PWN Warszawa:
381-385.
Szypuła J. 2000. Przewodnik do ćwiczeń z biologii ryb. Wydawnictwo AR-T Szczecin.
Tandyrak R., Parszuto K., Górniak D., Kośnik P. 2009. Hydrochemical properties, bacterioplank-
ton abundance and biomass in meromictic Lake Starodworskie in 2004. Oceanological and Hydro-
biological Studies XXXVIII 4: 128-133
Tandyrak R., Gawrońska H. 2009. The Influence of Urban Development on the Water Mass
Dynamics in the Lake. Polish Journal of Environmental Studies, Series of Monographs, Water Protec-
tion Systems in Agricultural and Industrial Regions - Selected Problems 3: 81-86, HARD
Wziątek B., Martyniak A., Kozłowski J., Osewski M., 2004. Wzrost siei w jeziorze Wigry przed
i po utworzeniu Wigierskiego Parku Narodowego. Rocz. Augustowsko-Suwalski IV: 131-136.
162
Irresponsible aquarists –
a threat to native ichthyofauna?
Mirosław Półgęsek, Paulina Hofsoe,
Bartosz Mysłowski
Unit of Aquaculture, West Pomeranian University of Technology

in Szczecin,
e-mail: miroslaw.polgesek@zut.edu.pl,
paulina.hofsoe@zut.edu.pl, bartosz.myslowski@zut.edu.pl
Abstract
Plants and animals which, when penetrating new habitats, dominate the native flora
and fauna as well as cause damage to the environment, are known as invasive alien spe-
cies. According to the European Environment Agency, alien species can be found in all
European ecosystems. For many years, we have heard reports on foreign species of exotic
fish observed in Polish waters. This phenomenon is largely shaped by the growing interest
in their amateur culture. Among the exotic fish species most frequently spotted in our wa-
ters are: pumpkinseed sunfish, Nile tilapia, pirapitinga, piranha, oscar and leopard pleco.
The cases of finding such fish in Polish waters, mentioned in this paper, point to a growing
problem of uncontrolled release of aquarium fish into aquatic ecosystems in our country.
This is a threat to the ichthyological biodiversity of our native water bodies.
Introduction
An alien species is the one which spreads outside its natural range of occurrence.
The main motivation for fish introduction has been: aquaculture, ‘improvement’ of the
natural species composition, game fishing, control of undesirable organisms, aquaristics
as well as fish escapes from fish culture facilities and incidental introduction of fish, e.g.
from ballast tanks of ships.
Plants and animals which enter new habitats can dominate the native flora and fau-
na and cause damage to the environment. These organisms are referred to as invasive
alien species. As the European Environment Agency states, alien species can be found in
all European ecosystems (http://eco.org.pl).
Until today, the only successful introduction of fish to Polish freshwaters has been
that of carp (Cyprinus carpio), while the other attempts at introducing such foreign spe-
cies as silver or bighead carp (Hypophthalmichthys molitrix and Aristichthys nobilis) or
grass carp (Ctenopharyngodon idella) have not fulfilled the expectations. An excellent
example is the introduction into European waters of brown bullhead (Ictalurus nebulo-
sus) (Annonimus 2007). Its natural range encompasses water bodies in North America
(the Mississippi and Missouri Rivers and their basins). It arrived in Europe, including
163
freshwaters in Poland, at the end of the 19th century. It raised hopes as a game and com-
mercial fish, owing to its size (up to 0.5 m), tasty meat and small environmental require-
ments. However, as it often happens to species imported from their natural range, the
fish has turned into a dwarf form, reaching just 25 cm in length. Thus, it no longer has
any commercial role, but has changed into “a fish weed”, which poses a threat to many
valuable species of our ichthyofauna by foraging on their spawn.
In the 1990s, some reports from anglers who caught exotic fish in Germany raised doubts
among people engaged in fisheries. The same approach could be seen when reports on the
presence of aquarium species such as guppy (Poecilia reticulata) and swordtail fish (Xipho-
phorus helleri) in warm waters of power plants in the Czech Republic were published (Keszka
et al. 2008). For some years now, we have heard about alien fish species in Polish waters. We
can expect to hear such reports more and more often. This situation is largely shaped by the
growing interest in keeping exotic fish at home. As with other hobbies, trends and fashions
in aquaristics change. Many aquarium hobbyists change their interests, which makes them
get rid of the species they have kept before in their aquaria. The easiest, but also the worst
way to do so is to release unwanted species to nearby water bodies.
Cases of reported alien species
Pumpkinseed sunfish Lepomis gibbosus (Linneaus 1758)
The original range of pumpkinseed sunfish is the east of North America. It was im-
ported to Europe in 1887 as an aquarium fish and to keep in garden ponds. In the Odra
River basin this species was spotted in 1927 (Kujawa 2008). It lives in shallow, sluggish
waters which become warm very quickly (Więcaszek et al. 2006). It can choose river
forks, small lakes, oxbow lakes and canals to which warm water is discharged from
power plants. Due to the preferred habitats, in West Pomerania, it has most often been
recorded in the Odra River and its tributaries and in the spent cooling water canal of the
Dolna Odra Power Plant, where the fish has found suitable conditions for reproduction
and now appears in large numbers (Ładyński 2002).
The pumpkinseed sunfish has a high raised back, laterally compressed body and small
length (up to 20 cm). The coloration is bluish, olive green in hue. The body is covered with
many spots. Additionally, it has a large black spot on the gill operculum and another one,
smaller and red in colour, directly behind the black one. It becomes sexually mature at the
age of 1-2 years and spawns from May to June. Eggs are laid to holes in a gravel bottom.
Larvae hatch after 3 days and are carefully protected by the male (Więcaszek et al. 2006).
This species is not a protected one. On the contrary, it is perceived as an undesirable
species. In places where it was introduced, it has been found to produce adverse effect
on the environment. When present in mass numbers, it can be a dangerous predator
feeding on larval stages and fish fry of native species (Kujawa 2008).
Nile tilapia Oreochromis niloticus (Linneaus 1758)
The natural range of this fish is Africa, the Nile River drainage up to the equator,
stretching northwards to Israel and westwards to the coast of the Atlantic Ocean. The
Nile tilapia has strongly laterally compressed body with a high back. The sides of the
body are silver in colour with dark stripes. During the mating season, the underside and
head of males become red and pink. Its maximum size is 60 cm long and 4 kg in weight
(Więcaszek et al. 2006). It inhabits stagnant or sluggish waters. It is tolerant to oxygen
deficit. Its staple diet consists of plants but it can also eat zooplankton and benthic or-
ganisms (http://www.iop.krakow.pl/gatunkiobce/).
164
Irresponsible aquarists – a threat to native ichthyofauna?
The first specimens of the fish arrived in Poland (the Polish Academy of Sciences Cen-
tre in Gołysz) in 1990 from the former Republic of Czechoslovakia. It was also cultured
in the RSD Nowe Czarnowo (Brylińska 2000). In recent years, tilapia has been recorded
in some water bodies in Poland, such as Konińskie Lake, the Ruda River and the Rybnicki
Reservoir. In the latter one, it is even used as a game fish and local angling organizations
stock the reservoir with this species every few years. Anglers have also reported that near
the power plant in Łęg-Kraków and in Nowa Huta, there is quite a large population of
Nile tilapia, which reproduces there (http://www.iop.krakow.pl/garunkiobce/).
Pirapitinga Piaractus brachypomus (Cuvier, 1818)
The natural range of occurrence of this fish is the freshwaters of the Amazon and
Orinoco Rivers and their basins in South America. This is a popular species to keep in
aquaria. Red-bellied pacu, as it is also known, has a high body (up to 47.5% of the body
length) which is laterally compressed. It grows to 70-80 cm long and lives up to 25 years.
The body is silver with a bright red belly (Więcaszek et al. 2006). Sometimes it has dark,
overlapping spots in the lower part of the body. Between the pectoral fins and the anal
opening it has a sharp keel. The fish feeds on insects and plants.
In Poland, there have been single reports of anglers who caught this species. In 2002,
it happened twice near Szczecin. The first case was on 20th June, when some pirapitingas
were captured in the spent cooling water canal of the Pomorzany Power Plant, and the
other one took place on 19th August, when a specimen of this species was captured in
Dąbie Lake (Więcaszek et al. 2007).
Introduction of the species Piaractus brachypomus could be a serious problem to the
Polish ichthyofauna, as this fish species is likely to be a source of parasites dangerous to
the fish living in the Polish watercourses. On the other hand, pirapitinga could be a valu-
able trophy for anglers.
Red piranha Pygocentrus nattereri (Knerr, 1858)
This species is a representative of the Characiformes and originates from South
America. It is broadly distributed in the basins of the Amazon, Orinoco and upper Parana
Rivers. This piranha has a silver body with numerous shiny speckles. The belly and the
underside of the neck are pink-red and the fins are dark. The coloration changes slightly
depending on the fish’s age and origin. In the wild, the fish can grow to 50 cm, but most
often do not exceed 30 cm in length (Więcaszek et al. 2007). This species lives in flocks,
which can gather up to tens of hundreds of individuals when waters become shallower.
They are distinctly meat-eating fish.
Due to their predatory nature, cases of catching piranha in our country are loud in
mass media. Tabloids often publish reports on people being bitten by this fish while
bathing in lakes. Such information, however, should be treated with utmost caution.
Reports on this fish being caught come from all over Poland, e.g. the Rybnicki Reservoir
or the Vistula River near Kraków.
Oscar Astronotus ocellatus (Agassiz, 1937)
This is a large species of the family Cichlidae, which naturally appears in South America, in
the basin of the Amazon River in Peru, Colombia, Brazil up to French Guyana. It grows to 45 cm
in length and has a strongly laterally compressed body. The coloration of this species is varied
(Kahl et al. 1997), most often it has a grey and olive green back with yellow, red and brownish
coating. Near the tail fin it has a characteristic spot in the shape of an eye on a peacock’s tail. It
is a voracious fish species and feeds on any kind of food it can swallow (Siudakiewicz 2007).
165
In 2007, one of the anglers’ online discussion groups posted a report on catching
an oscar fish, which had lived in a reservoir in Chojnów for three years (Anonymous
2007). The fish was in good body condition, which was made evident when it tried to
eat food. Another case described is a fish 26 cm long and weighing 410g, captured in
Grzymisławskie Lake in Śrem (http://www.haczyk.pl/forum/).
Leopard pleco Pterygoplichthys gibbiceps (Kner, 1854)
This species belongs to the family Loricariids (Loricariidae). It inhabits the basins of the
Amazon River and Orinoco in Venezuela, Brazil and Peru (Kahl et al. 1997). In the wild, it grows
to 50 cm in length. It has an abdominally compressed body and a characteristic suckermouth.
The basic coloration is brownish or orange-golden with dark dots scattered all over the body.
In the wild, this fish feeds on plants, most often scraping algae overgrowing stones.
The first case of catching a leopard pleco in Poland was in 2006, in the Brda River In
Bydgoszcz. The water temperature then was 7oC. This specimen was baited with a Den-
drobaena worm, and it weighed 0.7 kg (Keszka et al. 2008).
Summary
The above cases of catching exotic fish clearly show that the problem of uncontrolled re-
lease of fish will become more and more serious. This can create many dangers to the ichthyo-
logical biodiversity of our native water bodies. These alien fish often originate from tropical
parts of the world so the problem of their undesirable presence solves itself in winter, which
these fish cannot survive. However, when they enter waters near power plants, which are kept
warm, they have a chance of surviving winter and even of reproducing. Indigenous predators
will not necessarily be able to exterminate all unwanted fish. In that case, their population can
grow rapidly and uncontrollably, which may cause losses in the fish assemblages of native
species. But a more severe threat posed by alien fish is not connected with their growth and
development but with the risk that they can bring different parasites, fungi, bacteria, viruses or
pathogenic protozoans. The diseases caused by these pathogens can be extremely dangerous
to native fish, which lack resistance to this new pathogenic factor.
References
Annonimus 2007. Z wody obcej do polskiej. Magazyn Akwarium. 63: 6.
Brylińska M. (Ed.) 2000. Ryby słodkowodne Polski. Wydawnictwo Naukowe PWN, Warszawa.
Grabowska J., Witkowski A., Kotusz J. 2008. Inwazyjne gatunki ryb w polskich wodach -
zagrożenie dla rodzimej ichtiofauny. Nowa Rzeczywistość, PZW: 90-96.
Kahl W., Kahl B., Vogt D. 1997. Atlas ryb akwariowych. Vrglags-GmbH & Co Stuttgart 102, 200.
Keszka S., Panicz R., Tański A. 2008. First record of the Leopard pleco, Pterygoplichthys gib-
biceps (Actinopterygii, Loricariidae) in the Brda River in the centre of Bydgoszcz (Norhtern Poland).
Acta Ichtiol. Piscat. 38(2): 135-138.
Kujawa R. 2008. Bass słoneczny - ryba z temperamentem. Nasze Akwarium 102: 5-8.
Ładyński M. 2002. Bass słoneczny. Nasze Akwarium. 34: 16-19.
Siudakiewicz S. 2007. Pielęgnica pawiooka. Nasze Akwarium. 84: 28-30.
Więcaszek B., Keszka S., Antoszek J. 2007. Pirapitinga (Piaractus brachypomus Cuvier, 1818)
– an alien species in the natural water Bains within the Szczecin city area (NW Poland). In: Book of
abstracts XII European Congress of Ichthyology [Eds. Buj I., Zanella L., Mrakovcic M.], Wyd. Tipomat
doo, Zagreb: 20-21.
Więcaszek B., Krzykawski S., Keszka S., Antoszek A. 2006. Ryby w akwakulturze i akwaturystyce.
Wydawnictwo Akademii Rolniczej w Szczecinie.
http://www.haczyk.pl/forum/viewtopic.php?p=67727&sid=9c6dd8673b1e39be7d27c84428c25036
accessed on: 14.12.2010
http://www.iop.krakow.pl/gatunkiobce/default.asp?nazwa=default&je=pl accessed on: 14.12.2010
http://eko.org.pl/index_trendy.php?dzial=5&kat=25&art=1560 accessed on: 14.12.2010
166
Changes in the oxygen regime of
lakes and their effect on pelagic
ichthyofauna
Włodzimierz Marszelewski, Adam Solarczyk
Nicolas Copernicus University in Toruń, Faculty of Biology and

Earth Sciences, Department of Hydrology and Water Management,
ul. Gagarina 9, 87-100 Toruń,
e-mail: marszel@umk.pl, adamsol@doktorant.umk.pl
Abstract
The Polish ichthyological literature contains many papers on the effectiveness of ven-
dace management in lakes, most often evaluated on the basis of the morphometry of
lakes and frequency of vendace stock releases. Much rarer are examples of the research
on optimum habitats for this fish based on concentrations of water dissolved oxygen
and water temperature. Meanwhile, the environmental conditions, such as consider-
able decrease in oxygen conditions in lakes, especially since the 1970s and 1980s, seem
very important. The objective of this research has been to evaluate habitat conditions
for pelagic fish, taking as an example vendace in some lakes in the Brodnickie Lakeland
examined in 2007-2009. The lower borderline of a vendace habitat was assumed at 2
mgO2.dm-3 and the upper one – 18oC. Different rates and different ranges of the changes
in habitats of vendace have been observed both in the lakes during the subsequent
years and between the lakes in the same years. Attention has been paid to different time
periods when the temperature of water reached 18oC depending on the meteorological
conditions. The percentage of habitats of vendace to the whole volume of each analyzed
lake proved to be highly variable over the years. The results of our studies on the thermal
and oxygen conditions for the purpose of establishing boundaries of habitats of vendace
can be used in the future for evaluation of the effect of climatic changes on the extent
of pelagic ichythyofauna’s habitats and their parameters.
Introduction
Over the past few years, there has been an increasing number of studies on ther-
mal and oxygen conditions in lakes with the aim of evaluating habitats of pelagic fish.
This development is connected with a growing rate of eutrophication of lakes due to
their contamination and climatic changes. A habitat occupied by pelagic fish can be
167
established according to concentrations of oxygen and water temperature. The lower

boundary of such a habitat is limited by a low concentration of oxygen. For vendace
(Coregonus albula) it is set at 2 mgO2.dm-3 (Bernatowicz et al. 1975); for lake cisco
(Coregonus artedi) it is 3.1±1.3 mgO2.dm-3 (Aku et al. 1997) and for lake trout (Salveli-
nus namaycush) it is 4 mgO2.dm-3 (Clark et al. 2003). The upper boundary is established
by water temperature: 18-19oC for vendace (Dembiński 1971), 11.8 ±2.1°C for lake
cisco (Aku et al. 1997) and 10oC for lake trout (Clark et al. 2003). Determinations of
areas of fish habitats are helpful in running rational fish management. Hydroacoustic
tests have proven that a population of vendace can constitute nearly 100% of all the
fish in the hypolimnion of some lakes (Doroszczyk et al. 2007).
In the past years, more attention was paid to the effect of the morphometry of
lakes (especially average depth) on the effectiveness of vendace management (Leopold
et al. 1998) or to the frequency of vendace stock releases (Leopold and Wołos 1998).
Another important research area was the influence on some environmental param-
eters (the percent oxygen saturation of the hypolimnion, BOD5, total phosphorus Ptotal,
total nitrogen Ntotal, chlorophyll a, Secchi’s disc) on the effects of coregonid fish man-
agement (Bnińska and Wołos 1998). The role of oxygen conditions in lakes and the
fact that in most cases they are the factor which limits the presence of ichthyofauna
were noticed by Kubiak (2003), who emphasized that oxygenation of species which
require much oxygen, such as whitefish (Coregonus lavaretus) should be at least 70%.
However, attempts at delineating boundaries of optimum habitats for vendace ac-
cording to measured concentrations of dissolved oxygen and water temperature have
been rare. These parameters, for example, have been used as auxiliary ones during
the acoustic monitoring of vendace in the lakes Pluszne, Łańskie, Maróz and Mielno
(Doroszczyk et al. 2007).
The purpose of this study has been to determine habitat conditions of pelagic fish,
with vendace in three lakes in the Brodnickie Lakeland, i.e. Zbiczno, Łąkorz and Ciche
investigated in 2007-2009 taken as a case study. The oxygen and temperature condi-
tions were determined from the spring to the autumn circulations. The main part of
the paper is preceded by some general information on changes in oxygen conditions
in lakes lying in north-eastern Poland.
Changes in oxygen conditions in lakes of north-eastern Poland

Changes in the concentration of oxygen in many lakes since the mid-20th century are
presented below on the basis of mostly unpublished data from the 1960s, provided by
the Inland Fisheries Institute in Olsztyn, and the data from the turn of the 20th and 21st
centuries obtained from the Voivodeship Inspectorates for Environmental Protection as
well as the authors’ own studies. An analysis of this type of data for 349 lakes situated
in north-eastern Poland was accomplished by Marszelewski (2005). The results of that
analysis imply that the concentration of oxygen in all the lakes decreased considerably.
In the 1960s (and before) oxygen was present in most of the lakes in the whole vertical
column, although its amounts near the bottom were variable. Most oxygen near the
bottom (8-9 mg.dm-3) was found in polymictic lakes. In deep dimictic lakes, the con-
centration of oxygen near the bottom was lower, usually from 1 to 3 mg.dm-3. There
168
Changes in the oxygen regime of lakes and their effect on pelagic ichthyofauna
was one exception, i.e. Hańcza Lake, the deepest lake in the North European Plain, in
which the amount of oxygen above the bottom remained on a similar level of 9 to 10.5
mg.dm-3 throughout the whole 20th century. Quite good oxygen conditions have also
been sustained in the profundal zone of Wigry Lake, although in that case a decrease in
the concentration of oxygen, from 6 to 3.5 mg.dm-3, is evident.
Oxygen conditions at the turn of the 20th and 21st centuries looked completely dif-
ferent. In most dimict lakes, shortage of oxygen during summer seasons was recorded
not only in the deepest parts of lakes but also at a depth of just a few meters. Presence
of oxygen was most often limited to the epilimnion. Thus, in lakes with poor water cir-
culation and shallow epilimnion, oxygen was not present below 4 meters of depth. As a
consequence, a large decrease in the oxygenation of hypolimnions appeared, alongside
a diminished thickness of the water layer containing oxygen, which Marszelewski (2005)
determined quantitatively as the lake’s oxygenation index Nj (quotient of the thickness
of a water layer with oxygen to the maximum depth of the lake).
Some improvement of the oxygen conditions at the turn of the 20th and 21st cen-
turies has been observed in just a few cases, e.g. in Mamry and Ełckie Lakes. In Mamry
Lake, oxygenation of the hypolimnion rose from 21.8 to 33.4%, and in Ełckie Lake – from
10.6 to 26%, although in the deepest part of the latter lake, near the bottom, there were
anoxic conditions. Examples of changes in oxygen conditions in lakes are presented in
Table 1.
A large amount of data documenting oxygen conditions in lakes during the sum-
mer stagnation in the mid-20th century and then at the turn of the 20th and 21st cen-
turies enabled us to investigate their regional variation in north-eastern Poland (Tab.
2). In the mid-20th century, the average oxygenation of the hypolimnions of lakes was
the highest in the Suwałki Lakeland, where it reached 35.2%. Good oxygen conditions
with an average oxygenation of hypolimnions of about 20% appeared in seven lake
districts in the central and western parts of the region, including the Olsztyńskie and
Chełmińskie Lakeland as well as the Great Masurian Lakes (Tab. 2). The lowest oxy-
genation was found in the lakes of the Iławskie Lakeland (14%) and on the Masurian
Plain (11.9%).
The concentration of oxygen above the bottom in the lakes’ deepest areas changed
on a similar scale. In the earlier of the two compared time period, the average oxygen
concentration was from 3.4 mg.dm-3 in the Suwalskie Lakeland to 1.3-1.2 mg.dm-3 in
the Iławskie Lakeland and on the Masurian Plain. In the later years, it ranged from 1.6
mg.dm-3 in the Suwalskie Lakeland to 0.0 mg.dm-3 in the Chełmińskie Lakeland (Tab. 2).
The average concentration of oxygen above the bottom of the lakes in the whole area of
north-eastern Poland decreased from 1.8 to 0.4 mg.dm-3, which – when converted into
the water saturation with oxygen – corresponds to 15.1 to 2.9%.
Due to a large depletion of oxygen in dimict lakes, the range (depth) of oxygen pres-
ence, which can be determined by the value of a lake’s oxygenation index, has been
reduced. Its extreme values in the 1960s were from 84.3% in the lakes on the Masurian
Plain to 100% in the Ełckie Lakeland, but at the end of the 20th c., they fell to 29.6%
in the Chełmińskie Lakeland to 82.8% on the Augustowska Plain. For all the analyzed
dimictic lakes, the oxygenation index decreased by an average from 94.1 to 65.4%,
169
which should be considered as a very undesirable change in the habitat conditions for
fish (Fig. 1).
Table 1. Comparison of oxygen conditions in some dimict ic lakes in the 1960s (acc. to unpublished data col-
leted by the Inland Fisheries Institute in Olsztyn) and at the turn of the 20th and 21st centuries (Marszelewski
2005). Key: NTH – average hypolimnion oxygenation, ITD – amount of oxygen above the bottom, Nj – lake
oxygenation index (quotient of the thickness of water layer with oxygen to maximum depth of the lake).
KWJM – the Great Masurian Lakes.
NTH ITD Nj
Lake Lakeland % mg·dm-3 %
1960s 1990s 1960s 1990s 1960s 1990s
Ciche Brodnickie 28.4 0.0 3.0 0.0 100.0 60.0
Skarlińskie Brodnickie 55.0 0.0 5.8 0.0 100.0 73.0
Ilińsk Iławskie 25.3 0.0 2.8 0.0 100.0 22.0
Ruda Woda Iławskie 18.1 3.7 1.2 0.0 100.0 90.0
Dadaj Olsztyńskie 37.3 20.1 3.6 0.1 100.0 100.0
Luterskie Olsztyńskie 40.6 0.0 4.2 0.0 100.0 48.0
Narie Olsztyńskie 36.7 26.4 4.2 1.3 100.0 100.0
Tejstymy Olsztyńskie 63.5 4.3 6.0 0.6 100.0 100.0
Wukśniki Olsztyńskie 66.9 53.9 6.8 4.6 100.0 100.0
Czos Mragowskie 47.2 12.3 2.0 0.0 100.0 66.0
Juno Mrągowskie 23.1 0.0 2.0 0.0 100.0 18.0
Piłakno Mrągowskie 43.2 17.6 4.6 2.6 100.0 100.0
Dobrzyń KWJM 48.2 0.0 4.0 0.0 100.0 38.0
Mamry KWJM 21.8 33.4 0.9 1.5 100.0 100.0
Mikołajskie KWJM 0.8 0.0 0.0 0.0 62.0 46.0
Niegocin KWJM 2.5 0.0 0.3 0.0 100.0 15.0
Święcajty KWJM 5.3 3.3 0.0 0.0 75.0 61.0
Tałty-Ryńskie KWJM 16.5 0.0 1.4 0.0 100.0 27.5
Ełckie Ełckie 10.6 26.0 0.1 0.0 100.0 89.0
Łaśmiady Ełckie 49.9 0.0 4.6 0.0 100.0 21.0
Selmęt Duży Ełckie 55.1 5.0 5.4 0.5 100.0 100.0
Hańcza Suwalskie 80.1 82.5 8.8 9.7 100.0 100.0
Krzywe Wig. Suwalskie 19.4 0.0 2.0 0.0 100.0 25.0
Szelmęt Duży Suwalskie 48.7 36.8 4.4 3.1 100.0 100.0
Serwy August. Plain 42.3 29.4 3.0 2.0 100.0 100.0
Kownatki Masur. Plain 0.0 0.0 0.0 0.0 39.0 29.0
Omulew Masur. Plain 1.6 0.1 0.1 0.0 100.0 31.0
Roś Masur. Plain 32.5 0.0 2.1 0.0 100.0 28.0
170
Table 2. Changes in oxygen conditions in dimictic lakes during summer stagnation seasons according
to the physico-geographical regions of north-eastern Poland (Marszelewski 2005).
Key: Nj – lake oxygenation index (quotient of the thickness of water layer with oxygen
to maximum depth of the lake), KWJM – the Great Masurian Lakes.
Average oxy- Oxygen concentration Lake oxygen-
Lakeland genation of above bottom ation index
hypolimnion
(number of lakes; mg·dm-3 % Nj (%)
(%)
total surface area in ha)
1960s 1990s 1960s 1990s 1960s 1990s 1960s 1990s
Dobrzyńskie (22; 1715) 20.5 1.0 1.7 0.1 14.2 0.9 94.9 53.7
Chełmińskie (23; 1698) 22.4 0.0 1.6 0.0 14.4 0.0 92.6 29.6
Brodnickie (19; 2205) 21.4 1.9 1.8 0.1 15.5 0.7 92.9 42.4
Iławskie (39; 8189) 14.0 1.1 1.3 0.1 11.3 0.5 91.5 54.6
Garb Lubawski (7; 492) 18.7 3.6 1.4 0.5 12.1 4.1 95.2 71.8
Olsztyńskie (50; 12030) 21.8 6.9 2.0 0.5 16.9 3.9 92.4 62.7
Mrągowskie (41; 7746) 18.9 3.5 1.6 0.3 13.6 2.7 93.0 53.3
KWJM (34; 32337) 19.0 6.2 1.5 0.3 12.8 2.3 97.1 51.9
Ełckie (39; 9642) 22.1 4.7 1.9 0.2 16.8 1.3 100.0 57.5
Suwalskie (30; 5810) 35.2 20.0 3.4 1.6 27.9 12.4 96.0 70.8
Augustowska Plain (10; 1996) 21.5 8.1 1.9 0.6 15.0 5.3 100.0 82.8
Masurian Plain (26; 7042) 11.9 2.1 1.2 0.1 10.1 1.1 84.3 45.5
NE Poland (340; 90902) 20.6 4.9 1.8 0.4 15.1 2.9 94.1 56.4
A slightly less serious

deterioration of the oxygen
conditions above the lake
bottom appeared in the
group of polymictic lakes,
which was a result of their
morphometric features,
more favourable in this re-
spect. However, in some of Figure 1. Average oxygenation of dimictic (D) and polymictic (P) lakes
during the summer stagnation (Marszelewski 2005).
the polymictic lakes, lack of
oxygen in the benthic water layer and presence of sulphur hydrogen were recorded back
in the 1960s, for example, in Jeziorak Lake such conditions were documented at a depth
of 8-10 m in August 1963 (Jankowski 1966). This confirms the fact that shallower lakes, in
which water exchange relies on the influx and discharge of water from and to the drainage
basin (catchment), are more rapidly polluted. The average oxygen concentration in pol-
ymictic lakes in the mid-20th century, in particular regions, ranged from 3.5 to 7.5 mgO2.
dm-3, but at the end of the 20th century it varied from 5.5 to 0.0 mgO2.dm-3 (Tab. 3).
171
Table 3. Changes in oxygen conditions in polymictic lakes during summer stagnation seasons according
to the physico-geographical regions of north-eastern Poland (Marszelewski 2005). Key: Nj – lake oxygenation
index (quotient of the thickness of water layer with oxygen to maximum depth of the lake),
KWJM – the Great Masurian Lakes.
Oxygen concentration above bottom Lake oxygenation

Lakeland mg dm-3 % index Nj (%)
1960s 1990s 1960s 1990s 1960s 1990s
Dobrzyńskie 7.5 4.1 78.8 42.6 100.0 84.0
Chełmińskie 6.6 2.9 70.7 32.2 100.0 81.4
Brodnickie 3.5 0.0 36.5 0.0 82.4 50.5
Iławskie 5.6 5.5 59.4 59.7 96.0 97.3
Olsztyńskie 4.2 2.3 42.2 25.5 100.0 86.7
Mrągowskie 5.8 3.1 59.7 33.4 100.0 88.0
Great Masurian Lakes 4.6 3.0 49.2 32.5 94.6 86.3
Ełckie 4.0 2.8 41.8 28.9 100.0 76.2
Suwalskie 4.0 3.0 41.4 31.1 100.0 90.8
Masurian Plain 5.4 3.6 57.0 40.9 100.0 85.2
Urszulewska Plain 6.8 3.1 71.5 33.4 100.0 93.2
NE Poland 5.3 3.0 55.3 32.7 97.5 83.7
Intensive water circulation in polymictic lakes largely limits the appearance and sus-
tenance of anoxic conditions in deeper parts of these lakes. Thus, the decrease in the
oxygenation of water in these lakes was much smaller than in dimictic ones. In the
1960s, the average value of the oxygenation index in all polymictic lakes was 97.5% and
in the last decade of the 20th c. it fell to 83.7% (Fig. 1). The difference in the average
oxygenation of the lakes in the physico-geographical regions analyzed in this paper did
not usually exceed 15%.
The above data clearly indicate that in the 1970s and 1980s the oxygen conditions
in the lakes situated in north-eastern Poland drastically deteriorated. During the same
time, an evident decline in catches of vendace from 463 lakes in the same part of Poland
was observed (Skrzypczak and Mamcarz 2006). This decline has been attributed mainly
to changes in the fishery management. In the light of the available literature and the
authors’ own investigations, it seems that lower catches of this fish species were not
caused by economic reasons alone. Environmental factors, including oxygen conditions,
play an important role in the effectiveness of vendace management.
Determination of the optimum habitat for vendace (Coregonus albula)

An optimum habitat for vendace has been determined in three lakes in the Brodnickie
Lakeland: Zbiczno (182.9 ha), Łąkorz (161.8 ha) and Ciche (110.8 ha). They are ribbon
172
lakes, whose maximum (mean) depths are 41.3m (11.7m), 30.3m (11.6m) and 13.4m
(6.8m), respectively. As they are situated close to one another, it was presumed that are
subjected to similar meteorological conditions. The trophic state of these lakes, defined
acc. to Carlson (1977), was also similar, i.e. mesoeutrophic (Tab. 4). Zbiczno Lake and
Łąkorz Lake are vendace lakes while Ciche Lake is a bream lake. All the lakes have been
stocked with vendace by the Polish Angling Association, Region of Toruń. In 2001-2010,
the highest amount of fish fry was introduced to Zbiczno Lake (up to 41.9 thousand in-
div..ha-1) and the smallest one – to Ciche Lake (up to 16.2 thousand indiv..ha-1). In all the
lakes, the smallest amounts of vendace stock material were introduced in 2010 (acc. to
unpublished materials of the PAA, Region in Toruń). The catches of vendace in these lakes
were low and variable. In some years, no catches of this fish species were performed in
Ciche and Łąkorz Lakes (Tab. 5). Catches of vendace in Zbiczno Lake – compared to the
amounts of fish fry released – were also low (up to 2.19 kg.ha-1 in 2005). Meanwhile, with
the average amount of stock material released in 2001-2010, such as from 18.8 thousand
kg.ha-1 (Zbiczno Lake) to 2.9 thousand kg.ha-1 (Ciche Lake) (Fig. 2), the expected catches of
vendace should be much higher, reaching – according to the model elaborated by Leopold
et al. (1998) – 14.36 kg.ha-1 in Zbiczno Lake, 7.85 kg.ha-1 in Łąkorz Lake and 2.44 kg.ha-1
in Ciche Lake. However, it should be added that the above model was worked out on the
basis of just two parameters: the mean depth of a lake and amounts of stock material re-
leased. Similar results of the effectiveness of vendace catches were obtained from another
model, which includes the frequency of fish fry releases (Leopold and Wołos). They were:
14.38 kg.ha-1 in Zbiczno Lake, 7.35 kg.ha-1 in Łąkorz Lake and 3.72 kg.ha-1 in Ciche Lake.
In 2005-2008, catches of vendace compared to the total fishery catches were highly
varied in the analyzed lakes and between the years. The highest percentage of vendace
in the total catches was recorded in Zbiczno Lake (from 35% in 2005 to 4% in 2008),
and the lowest one – in Ciche Lake (from 2.5% in 2007 to 0% in 2005). Noteworthy is
the fact that catches of vendace in Zbiczno Lake, despite regular fish fry releases (Fig.
2), evidently tended to de-
crease. At this stage of our
research, it is difficult to
state firmly whether this ef-
fect is mainly caused by en-
vironmental changes or the
way the lake is managed.
In all the three lakes, fish-
ery catches are larger than
angling ones. This can be
concluded from the data of
2007 found in the report pre-
pared by Wołos et al. (2009).
The total catches by anglers Figure 2. Volume of released (thousand of indiv. ha-1) vendace stock
in all the three lakes are dom- material into Zbiczno (A), Łąkorz (B) i Ciche (C) Lakes in 2001-2010
(acc. to the data of the PAA, Region in Toruń) key: 1 – the average value
inated by cyprinids (Tab. 6). of the years of fish fry releases.
173
Table 4. Indices of the trophic state of the lakes during the summer stagnation (acc. to unpublished data from
the Voivodeship Inspectorates for Environmental protection in Bydgoszcz and Olsztyn). Key: SD- Secchi’s disc,
Chl. a – chlorophyll a, TP – total phosphorus.
Trophic indices TSI (Carlson 1977)

Lake
SD (m) Chl. a (mg m ) -3
TP (mg m ) -3
SD Chl. a TP
Zbiczno (2009) 2.1 4.68 19 49 46 47
Łąkorz (2003) 4.4 7.30 40 39 50 57
Ciche (2008) 2.2 7.15 26 49 50 51
Table 5. Catches of vendace (kg ha-1) in 2005-2008

(acc. to the data of the Polish Angling Association, Region in Toruń).
Year Zbiczno Lake Łąkorz Lake Ciche Lake

2001 1.10 0.00 0.00
2002 0.55 0.31 0.00
2003 0.82 0.00 0.00
2004 0.45 0.08 0.00
2005 2.19 0.10 0.00
2006 1.34 0.00 0.20
2007 0.23 0.01 0.36
2008 0.21 0.39 0.08
2009 0.39 0.05 0.00
2010 0.02 0.00 0.00
Table 6. Angling catches (kg ha-1) in 2007 acc. to registers (Wołos et al. 2009).
Fish species Zbiczno Lake Łąkorz Lake Ciche Lake

Bream 0.77 0.09 0.20
Tench 0.08 0.00 0.05
Pike 0.51 0.47 0.58
Eel - 0.01 -
Perch 0.05 0.10 0.08
Roach 1.19 0.77 0.75
Others 0.05 - 0.02
Total catches 2.65 1.44 1.68
This study contained an attempt to determine a suitable habitat for vendace includ-
ing the aspect of oxygen and thermal conditions. From April to December 2007, 2008
and 2009, the vertical temperature and water dissolved oxygen distribution in the deep-
est parts of the lakes was examined every two weeks (in 2008 and 2009) or every month
(in 2007). In order to facilitate comparison of the results obtained in each year, the meas-
urements were made on approximately the same days (±3 days). The optimum habitat
conditions for vendace were established according to the methodology suggested for
174
lake trout by the Ontario Ministry of the Environment and the Ontario Ministry of Natu-
ral Resources (1993) and by Dillon et al. (2003). The lower habitat boundary was set at
2 mgO2.dm-3 (Bernatowicz et al. 1975), and the upper one – at 18oC (Dembiński 1971).
The results of our investigations are presented in Figure 3. They reveal differences in the
rate and extent of the environmental changes affecting vendace, both in the same lake
in subsequent years and in the same year between the lakes.
In all the three lakes and in most of the cases, the upper (thermal) boundary of the
habitat appeared first. This was associated with an increase in the water temperature
above 18oC. Such temperature was recorded from late May 2007 and 2008, but nearly
a month later in 2009, which was caused by a long and cool spring. The thermal limit
lasted until mid-August or mid-September and, in each year, depended on the meteoro-
logical conditions and the mictic features of the lakes (Fig. 3).
In the summer season, the water temperature of 18oC, in most cases, was recorded
near the upper boundary of the metalimnion. However, it appeared on different dates.
Due to the low air temperature during the spring and early summer in 2009, such water
temperature was not recorded until the turn of June and July (Fig. 4). As a result, the
upper boundary of vendace habitats appeared later than in the preceding years.
The lower (oxygen) boundary of vendace habitats appeared much later, especially in
Zbiczno and Łąkorz Lakes. The formation of that boundary was due to the depletion of
oxygen caused, for example, by mineralization of organic matter in benthic sediments and
a decrease in the concentration of oxygen concentration below 2 mgO2.dm-3. In Zbiczno
Lake, this oxygen limitation appeared as late as mid-July in 2007 and 2008 and mid-June
2009 (due to a relatively short period of spring homothermy and consequently poor oxy-
genation of the deepest water layers). Slightly sooner, at the end of June, the lower bound-
Figure 3. Variability in the boundaries of vendace habitats (% of the total volume of the lake)
in 2007-2009 in Zbiczno (A), Łąkorz (B) i Ciche (C) Lakes.
175
ary of the habitat appeared

in Łąkorz Lake. The same sit-
uation was observed in each
year. The first to have the
oxygen-related boundary of
a habitat suitable for ven-
dace was Ciche Lake, where
it occurred in early May. In
this lake, which is shallower
and holds less water, only
some of the sedimenting or-
ganic matter is mineralized
in the water column, which
– compared to the other
two, deeper lakes – causes
greater accumulation of
organic substances in the Figure 4. Temporal and spatial variability in the extent of the metalimnion
in 2007-2009 in Zbiczno (A), Łąkorz (B) i Ciche (C) Lakes. Key: 1 – range
benthic sediments and thus of the metalimnion, 2 – water temperature 18oC.
a more rapid consumption
of oxygen. The oxygen-related limitation of the vendace habitats ended when the autumn
circulation began. This circulation started the earliest in Ciche Lake (mid-October) and the
latest in Zbiczno Lake (mid-December). These differences are mainly caused by the differ-
ences in the morphometric and mictic properties of the lakes (Fig. 3).
The percentage of vendace habitats in the total volume of the lakes was highly variable
over the years (Fig. 5). The maximum share of such habitats (100%) occurred during the
spring and autumn circulations. The minimum share appeared, in the vast majority of cases,
during the peak summer stagnation (mid-August ±5 days), when it fell to nearly 0%. Zbic-
zno Lake was an exception because there the minimum share of such a habitat was about
40% in 2007 and 2008, but in 2008 some delay, by nearly a month, in the occurrence of
the minimum percentage of that habitat was observed. In Zbiczne Lake, the percentage of a
vendace habitat close to 0 occurred only in 2009, which – as mentioned earlier – was caused
by specific hydro-meteorological conditions during a very short spring season.
Final remarks
This paper deals with the determination of habitat conditions for vendace in three lakes
stocked with this species, two of which are classified as vendace type of lakes. The lakes were
chosen for the study rather by chance, as they all belong to a lager group of lakes analyzed in
terms of the differentiation of oxygen conditions. The three lakes are classified as small ones,
but characterized by quite big differences in their morphometry. Therefore, we can hope
to have obtained quite interesting results, which point to high dynamics in the changes of
environmental conditions for vendace, both in the course of a year and during a longer time
period. Significant differences were observed in the habitats found in the analyzed lakes.
Small catches of vendace, compared to the expected ones resulting for the application of the
model elaborated by Leopold et al. (1998), may suggest that the habitat conditions in these
176
lakes are not suitable for the

growth and development
of this species. In Łąkorz
and Ciche Lakes, habitats of
vendace are greatly limited
in August, as they comprise
just a water layer between
the epilimnion and metalim-
nion. The fact that such en-
vironmental conditions force
vendace to dwell in that wa-
ter layer during the daytime
exposes the fish to pressure
exerted by predatory fish. It
does not seem possible that
cormorants (Phalacrocorax
carbo), which are not im-
plied as an actual threat to
vendace, could play an im-
portant role in reducing their
populations (e.g. Engström
2001, Gmitrzuk 2004, Krzy-
wosz 2008).
Detailed examination of
thermal and oxygen condi-
tions, for the sake of deter-
mination of habitats suitable
for vendace, can be used in
the future to assess the ef-
fect of climatic changes on
the range of habitats colo-
nized by pelagic ichthyofau-
na and their parameters. The
first such attempt has been
made (Elliot and Bell 2010).
The methodology used
for delineation of fish habi-
tats, presented in this pa-
per, can be also applied to Figure 5. Percentage of vendace habitats (%) to the total volume
determination of habitats of the lakes in 2007–2009.
of another valuable pelagic
fish, living in Poland, such as lake trout (Salmo trutta morpha lacustric). Studies on the
range of occurrence of lake trout based on its catches at different depths were conduct-
ed by Radtke (1999), although the actual habitat conditions were not determined.
177
Acknowledgement
The investigations were financed from the grant of the Polish Ministry of Science and
Higher Education no. N306 354836.
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178
Habitat conditions for the
ichthyofauna in the largest
lakes in the West Pomerania
Lake District
Jacek Kubiak, Sylwia Machula
West Pomeranian University of Technology, Department of Hydro-

chemistry and Water Protection, e-mail: jacek.kubiak@zut.edu.pl
Abstract
In 1970-2020, 13 largest lakes (>100 ha) in West Pomerania were investigated in re-
spect of the habitat conditions of ichthyofauna under progressing eutrophication. It was
found out that the oxygen conditions limited the presence of ichthyofauna, especially
in the lakes of high trophy level and in the waters lying in the hypolimnion and in the
benthic layers. Such properties as the content of suspension, water pH, concentration
of CO2 or undissociated ammonia did not limit the occurrence of fish. The best habitat
conditions for fish appeared in Ińsko Duże and Miedwie Lakes.
Introduction
The current state of lake eco-geosystems is mainly a consequence of progressing
eutrophication, perceived as changes caused by uncontrollable or insufficiently control-
led influx of biogenic substances (nitrogen and phosphorus) from external sources, or
due to the anthropogenic direct intervention into the internal structure of an ecosystem
(Lossow 1995, Zdanowski 1996, Kalff 2002). Phosphorus plays a particular role in this
process as it is the biogene held responsible for the level of trophy (Kalff 2002). Likewise,
excess organic matter, either allo- or autochthonous in origin, intensifies eutrophication
because mineral forms of biogenic substances are released during its mineralization
(Kalff 2002). At present, it is assumed that eutrophication is the most severe threat to
biodiversity of freshwater habitats (Hillbricht-Ilkowska 1998).
A variety of functions assigned to lakes and their great importance forces people to
undertake protection and reclamation measures in order to reverse or at least to halt the un-
desirable changes in these eco-geosystems, caused by their increased trophy (Lossow 1995).
Such actions will make it possible to run rational fishery management (Zdanowski 1993).
179
In overfertilized lakes, water is less suitable for municipal or industrial use, for fisher-
ies or for water recreation. Furthermore, lakes are an important link in water retention,
and as the water quality deteriorates and groundwater resources are becoming deplet-
ed, lakes turn into the major source of water of adequate quality, thus their role is ever
more important (Kubiak 2003).
The state of lakes in Poland is like that in most European countries. Over the past
years, more than 85% of lakes were characterized by the concentration of total phos-
phorus in water and water transparency typical for eutrophic water bodies (Hillbricht-
Ilkowska 1998, Lossow 1995). Recently, the level of trophy has been observed to decline
in some lakes (Kubiak et al. 2009).
In Poland, among the lakes covering more than 50 ha (1,026), just 12.6% are prog-
nosticated to attain the set environmental target, i.e. to have good quality water, in 2015
(Cabaj 2010). Considering the importance of lakes for the fisheries in West Pomerania,
the objective of this study has been to determine the habitat conditions for ichthyofauna
in the largest (over 100 ha) stratified lakes in this region, the conditions that have been
shaped under increased eutrophication.
Location of the examined lakes.

The examined lakes lie in the West Pomerania Lake District, which covers 9.7 thou-
sand km2 and is divided into 7 mesoregions, among which the Myśliborskie and Ińskie
Lake Districts, including the analyzed lakes, are distinguishable.
The Myśliborskie Lake District covers an area of about 1,810 km2; in its western part
(the chojeńsko-moryński complex) lie such lakes as Morzycko, Narost (both in the drain-
age basin of the Słubia River) and Jelenin (in the drainage basin of the Rurzyca River).
In the eastern part of the district (in the myśliborsko-lipiański complex), in the drainage
basin of the Myśla River, there are the lakes: Wądół (Lipiańskie Północne), Będzin, Chłop
Duży and Chłop Mały (Kondracki 2000).
The Ińskie Lake District is a mesoregion about 750 km2 in area; the following lakes
situated in this area were analyzed: Ińsko Duże, Ińsko Małe (Odnoga Linowska), Wiso-
la (Wisala, Stubnica) and Krzemień (in the drainage basin of the upper Ina River) and
Woświn Lake (in the drainage basin of the Ukleja River, a left tributary of the Rega River)
(Kondracki 2000).
Miedwie Lake, one of the largest lakes in the West Pomerania Lake District, lies on the
Pyrzycka Lowland, in the drainage basin of the Płonia River, about 8-10 km south-west
of Stargard Szczeciński (Cabaj 2010).
The analyzed lakes lie in an area which in the past was dominated by the so-called
large-scale agricultural production, characterized by commercial animal breeding and
application of large doses of artificial fertilizers (Kubiak 2003). This had a considerable
influence on the transport to these lakes of large loads of biogenes and a very intensive
process of eutrophication (Kubiak 2003, Cabaj 2010).
The geographical location as well as the trophic state and fisheries specification of
the lakes can be found in Table 1.
180
Habitat conditions for the ichthyofauna in the largest lakes in the West Pomerania Lake District
Table 1. Geographical location as well as trophic and fisheries characteristics of the analyzed lakes
(Kubiak 2003, Cabaj 2010).
Geo- Influ- Rate
Tolerance
graphi- Trophic Water ence of of eu- Lake fishery
Lake to degra-
cal loca- state mixing drainage trophi- type
dation
tion basin cation
52o59’02 N Hypereu- Moder-
Będzin Tachymictic Low Fast Pike-perch
14o56’28 E trophic ate
Chłop 52o59’47 N Mesoeu-
Eumictic High Small Slow Vendace
Duży 14o53’50 E trophic
Chłop 53o00’36 N
Eutrophic Eumictic High Small Slow Bream
Mały 14o54’28 E
Ińsko 53o26’35 N Mes- Eumctic/ Moder- Moder-

Moderate Vendace
Duże 15o32’36 E otrophic bradymictic ate ate
Ińsko 53o25’52 N Moder-
Eutrophic Tachymictic Moderate Fast Bream
Małe 15o30’59 E ate
52o54;48 N Mesoeu-
Jelenin Eumictic Moderate Small Slow Bream
14o30’06 E trophic
53o22’27 N Mesoeu- Moder- Moder-
Krzemień Eumictic Moderate Bream
15o32’17 E trophic ate ate
53o16’11 N
Mes-
otrophic/ Eumictic/ Moder- Moder-
Miedwie High Vendace
Mesoeu- bradymictic ate ate
14o52’59 E
trophic
52o5153 N Mesoeu- Bradymictic/ Moder- Moder-
Morzycko High Vendace
14o24,36 E trophic Eumictic ate ate
52o52’59 N Moder- Moder-
Narost Eutrophic Tachymictic Moderate Bream
14o30’20 E ate ate
53o00’38 N Hypereu-
Wądół Tachymictic Low Big Fast Pike-perch
14o57’17 E trophic
53o24’51 N Moder-
Wisala Eutrophic Tachymictic Moderate Fast Bream
15o33’04 E ate
53o31’34 N Mesoeu- Moder- Moder-
Woświn Eumictic Moderate Bream
15o25’20 E trophic ate ate

The habitat conditions for the fish in the analyzed lakes were assessed according to the
indices which are cited in the references as the most important for the physiology of fish.
Oxygen conditions and pH, which determine the normal course of life functions of fish
and their tolerance to unfavourable conditions of the external environment (Łukjanienko
1974, Opuszyński 1983), were analyzed. Other factors considered were the concentration
of CO2 dissolved in water, which shapes the effectiveness of gas exchange between a fish
organism and the environment, and NH3 concentration, which determines the efficiency
of the removal of NH3 from the circulatory system of fish through gills (Schmidt-Nielsen
181
1992). These indicators are largely conditioned by the transformation of matter in lakes,
which is especially intensive in water bodies of advanced trophy.
The paper relies on the authors’ own investigations (dating back to 1971-1985 and
1992-2010) and some unpublished materials produced in 1986-1991 at the Department
of Hydrochemistry and Water Protection of the University of Agriculture in Szczecin.
Some of the older data (1974-1975) regarding the lakes Mokrzycko, Będzin and Wądół
were found in the materials published by the Inspectorate for Environmental Protection
(Atlas 1977), and the ones concerning the years 1970-1971 about the lakes Ińsko Duże,
Ińsko Małe and Wisala were obtained the doctoral thesis of Nguyen (1972) while the
information about Miedwie Lake comes from the PhD thesis of Cabaj (2010).
Field methods
Studies on particular lakes were usually carried out in 5-year cycles. They included
at least the period of spring homothermy and summer stagnation. The timetable of
the studies is presented in Table 2. Samples of water were taken with the Ruttner ba-
thometer from the deepest sites of each lake, simultaneously measuring its temperature
(a mercury thermometer ±0.1şC). Content of oxygen and temperature were determined
along the vertical cross-section at every 1 meter. For the other analyses, in each of the lakes
during the peak summer stagnation, samples of water were taken from surface waters (0.5
m under the surface), at the border of the epilimnion and thermocline, the thermocline
and hypolimnion and from benthic waters (0.5 m above the bottom). Besides, two water
samples, uniformly spaced along the vertical cross-section, were collected from each water
layer. In the other limnological seasons, at their peaks, water sampling was always carried
out at the same depths, uniformly spaced along the vertical cross-section. Then, five sam-
ples were collected from lakes of the maximum depth less than 20 m and eight samples
from deeper lakes, including the surface and benthic layers. Only some of the results have
been used for writing this publication.
Laboratory methods
The water samples underwent determinations, with the generally adapted methods,
of the following indicators necessary for our evaluation of the habitat conditions for
fish:
• ammonia nitrogen – by direct nesslerization or with indophenol blue,
• amount of seston’s dry matter (total suspension) – by weight,
• oxygen – with Winkler’s method or with an oxygen meter manufactured by the
WTW company
• water reaction (pH) – conductometrically,
• CO2 dissolved with 0.05 NaOH,
• and the amount of undissociated ammonia according to Alabaster and Lloyd
(1980).
Spectrophotometers UV-Vis: Beckman-25 (Beckman) and Lambda-10 (Perkin Elemer)
were applied to colorimetric examinations. The timetable of the determinations is in-
cluded in Table 2.
182
Table 2. Timetable of the field tests. W – spring, L – summer, J – autumn, Z – winter;

figures after the letters stand for the number of tests in a given season of the year.
LAKE 1977 1980 1983 1985 1990 1993 1995 1996 1998 2005 - -
W-1; W-1; W-1; W-2; L- W-1; L- W-4; L- W-4; L-
L-2; L-2; W-1; W-1; W-1;
Będzin L-2;
L-2; J- 2; J-1;
L-2
1; J-1;
L-2
2; J-2; 2; J-2; - -
J-1;Z-1 J-1;Z-1 1;Z-1 Z-1 Z-1 Z-2 Z-2
1977 1980 1983 1985 1990 1993 1995 1996 1999 2005 - -
Chłop W-1; L- W-1; L- W-1; L- W-2; L- W-1; L- W-4; L- W-4; L-
Duży W-1; W-1; W-1;
2; J-1; 2; J-1; 2; J-1; 2; J-1; 2; J-1; 2; J-2; 2; J-2; - -
L-2; L-2 L-2
Z-1 Z-1 Z-1 Z-1 Z-1 Z-2 Z-2
1977 1980 1983 1985 1990 1993 1995 1996 1998 2005 - -
Chłop W-1; W-1; W-1; W-2; W-1; W-4; W-4;
Mały W-1; W-1; W-1;
L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- - -
L-2; L-2 L-2
1;Z-1 1;Z-1 1;Z-1 1;Z-1 1;Z-1 2;Z-2 2;Z-2
1970 1971 1978 1982 1987 1992 1996 1997 1998 1999 2003 2007
Ińsko W-1; W-1; W-1; W-1; W-1; W-4; W-4; W-4;
Duże W-1; W-1; W-1; W-1;
L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J-
L-2; L-2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 1;Z-1 1;Z-1 2;Z-2 2;Z-2 2;Z-2
1970 1971 1978 1982 1987 1992 1996 1997 1998 1999 2003 2007
Ińsko W-1; W-1; W-1; W-1; W-1; W-4; W-4; W-4;
Małe W-1; W-1; W-1; W-1;
L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J-
L-2; L-2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 1;Z-1 1;Z-1 2;Z-2 2;Z-2 2;Z-2
1979 1983 1988 1991 1995 1996 1999 - - - - -
Jelenin W-1; W-1; W-1; W-4;
W-1; W-1; W-1;
L-2; J- L-1; J- L-1; J- L-2; J- - - - - -
L-2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 2;Z-2
1978 1982 1987 1992 1996 1997 1998 1999 2000 2003 2007 -
Krzemień W-1; W-1;
W-1;
W-1;
W-4;
W-1; W-1;
W-4; W-1; L- W-4; W-4;
L-2; J- L-2; J- L-2; J- L-2; J- 1; J-1; L-2; J- L-2; J- -
L-2; L-2; L-2; L-2;
1;Z-1 1;Z-1 2;Z-2 2;Z-2 Z-1 2;Z-2 2;Z-2
1995 1987 1990 1993 1995 1997 1999 2001 2003 2005 2006 2007
Miedwie W-1;
W-1; W-1; W-1; W-1; W-1; W-1; W-1; W-1; W-1; W-1; W-1; L-
L-1; J-
L-2; L-2; L-2; L-2; L-2; L-2; L-2; L-2; L-2; L-2; 1; J-1
1;Z-1
1974 1979 1983 1988 1991 1995 1996 1999 2003 2008 - -
Morzy- W-1; W-1; W-1; W-1; W-4; W-4; W-4;
cko W-1; W-1; W-1;
L-2; J- L-1; J- L-1; J- L-1; J- L-2; J- L-2; J- L-2; J- - -
L-2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 1;Z-1 2;Z-2 2;Z-2 2;Z-2
1979 1983 1988 1991 1995 1996 1999 - - - - -
Narost W-1; W-1; W-1; W-4;
W-1; W-1; W-1;
L-2; J- L-1; J- L-1; J- L-2; J- - - - - -
L-2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 2;Z-2
1977 1980 1983 1985 1990 1993 1995 1996 1998 2005 - -
Wądół W-1; W-1; W-1; W-2; W-1; W-4; W-4;
W-1; W-1; W-1;
L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- - -
L-2; L-2 L-2
1;Z-1 1;Z-1 1;Z-1 1;Z-1 1;Z-1 2;Z-2 2;Z-2
1970 1971 1978 1982 1987 1992 1996 1997 1998 1999 2003 2007
Wisola W-1; W-1; W-1; W-1; W-1; W-4; L- W-4; W-4;
W-1; W-1;L- W-1; W-1;
L-2; J- L-2; J- L-2; J- L-2; J- L-2; J- 2; J-2; L-2; J- L-2; J-
L-2; 2; L-2; L-2;
1;Z-1 1;Z-1 1;Z-1 1;Z-1 1;Z-1 Z-2 2;Z-2 2;Z-2
1973 1982 1987 1992 1994 1995 1996 1997 1999 - - -
Woświn W-1; L- W-1; L- W-4; -
W-1; W-1; W-1; W-1; W-1; W-1;
2; J-1; 2; J-1; L-2; J- - -
L-2; L-2; L-2; L-2; L-2; L-2;
Z-1 Z-1 2;Z-2
183
Discussion of the results

Below we shall discuss the hydrochemical indices taken in this paper to evaluate the
habitat conditions for fish shaped in the examined lakes. Particular attention will be paid to
the extreme values of these indices, in respect of their limiting influence on the occurrence
of fish (Tab. 3).
The amount of ammonia dissolved in the waters of the analyzed lakes was varied. Typi-
cally, higher values of this index were determined for surface waters in summer than in
deeper layers in colder seasons of the year. Throughout the whole period of our studies, over
90% of all the results were within the range 0.000-0.010 mg NH3.dm-3, and just 2% of the
results (obtained mainly for Wądół and Będzin Lakes) were above 0.025 mg NH3.dm-3. This
value was never exceeded for the water samples taken from Chłop Mały and Chłop Duży,
Ińsko Duże and Ińsko Małe, Jelenin, Krzemień and Morzycko Lakes. The highest concentra-
tions of ammonia were observed in summer in surface waters of the lakes Wądół (0.076 mg
NH3·dm-3), Narost (0.071 mg NH3·dm-3) and Będzin (0.0590 mg NH3·dm-3). The average
value of this index in all the lakes was 0.0098 in epilimnion, 0.0048 in thermocline and
0.0029 mg NH3·dm-3 in hypolimnion. The lakes Ińsko Duże, Chłop Duży, Krzemień and
Jelenin were characterized by the lowest concentrations of ammonia – in surface waters
they were on average 0.007, 0.004, 0,002 and 0.004 NH3·dm-3, respectively (Tab. 3).
The water reaction (pH) in the analyzed lakes, like ammonia, was varied in time be-
tween particular lakes as well as along the vertical cross-section in each lake. As a rule,
this index was higher in summer in surface waters than in deeper water layers in colder
seasons. Throughout the whole period of investigations, over 88% of the measurements
of water pH were within the range of 7.0-8.5. The highest reaction (pH) was typical for
the surface waters of Wądół Lake, where this index would always assume the values
above 8.75 and on a few occasions rose above 9.00. Similarly, in Ińsko Małe, Wisala and
Będzin Lakes the value of pH was found to have exceeded that value. The above lakes
were also characterized by the highest variability of water reaction, both in surface and
benthic water layers as well as between these layers. Typically, the water reaction in
benthic waters was 0.4-0.6 lower and less variable. In none of the analyzed lakes, the
water pH fell below 6.50 and the lowest values of pH were determined in benthic waters
of the lakes Narost, Ińsko Małe and Wądół. The remaining lakes were characterized by
a distinctly lower water reaction, and its smallest fluctuations in the surface waters were
determined in Miedwie, Jelenin, Chłop Duży and Morzycko Lakes; in turn, the benthic
waters of Miedwie, Ińsko Duże, Jelenin and Wisala were the least variable in the values
of pH (Tab. 3).
Carbon dioxide dissolved in water appeared in the analyzed lakes within the range of 0.0
to 42.9 mg.dm-3. Higher concentrations of carbon dioxide were always found in benthic water
and in winter than in surface layers or in summer. In the surface waters, the average content
of CO2 varied from 5.3 (in Ińsko Małe Lake) to 9.5 mg.dm-3 (in Będzin Lake) and the maximum
range of variability was 0.0-27.2 mg.dm-3 (in Będzin Lake). A similar range of changes of this
index was recorded in Chłop Mały Lake (0.0-21.7 mg.dm-3). In the other lakes, the average value
of dissolved CO2 in the surface layer of water did not exceed 12.0 mg.dm-3. In the benthic layers,
this index was always higher than in the surface water, e.g. it exceeded 40 mg.dm-3 in Będzin
and Chłop Mały Lakes; in the other lakes, it was below twenty milligrams per litter at the most.
184
Table 3. Values of some hydrochemical indices which determine the habitat conditions for fish in the examined lakes.
Water Total
Lake O2 CO2 NH3 pH
layer suspension
surf. 10.9 (5.5-18.8) 21.4 (8.0-38.0) 9.5 (0.0-27.2) 0.011 (0.000-0.059) 7.92 (7.01-9.03)
Będzin
bottom 4.7 (0.0-12.9) 13.5 (7.0-27.0) 29.6 (7.8-42.9) 0.004 (0.000-0.030) 7.43 (6.79-7.95)
Chłop surf. 11.5 (8.2-13.2) 18.0 (10.0-32.0) 7.7 (0.0-9.80) 0.004 (0.000-0.014) 7.85 (7.50-8.68)
Duży bottom 5.9 (0.3-12.3) 8.7 (7.0-18.0) 8.8 (6.6-20.7) 0.001 (0.000-0.017) 7.45 (6.95-8.28)
Chłop surf. 11.7 (9.9-13.7) 14.7 (8.0-28.0) 6.4 (0.0-21.7) 0.002 (0.000-0.024) 8.00 (7.30-8.76)
Mały bottom 6.4 (0.0-11.1) 8.0 (5.0-12.0) 18.6 (2.8-41.8) 0.003 (0.000-0.014) 7.39 (6.70-8.19)
Ińsko surf. 11.5 (10.0-14.0) 13.8 (5.0-24.0) 6.0 (0.0-12.1) 0.007 (0.003-0.020) 7.79 (7.20-8.73)
Duże bottom 7.3 (3.2-11.9) 7.7 (5.0-14.0) 7.2 (2.0-19.8) 0.001 (0.000-0.003) 7.53 (7.10-7.93)
Ińsko surf. 11.0 (8.5-12.5) 12.0 (5.0-25.5) 5.3 (0.0-11.2) 0.003 (0.000-0.025) 7.89 (7.10-9.01)
Małe bottom 8.1 (0.0-11.6) 6.3 (4.0-11.0) 5.9 (4.0-21.0) 0.002 (0.000-0.007) 7.40 (6.56-7.88)
surf. 11.0 (8.8-13.0) 13.7 (4.5-28) 8.1 (4.0-11.0) 0.004 (0.000-0.024) 7.89 (7.68-8.70)
Jelenin
bottom 4.0 (0.0-10.2) 5.9 (3.0-11.5) 9.2 (4.0-14.0) 0.002 (0.000-0.008) 7.56 (7.23-8.01)
surf. 11.3 (9.3-14.0) 8.1 (3.0-19.5) 8.1 (0.0-9.5) 0.002 (0.000-0.016) 7.89 (7.50-8.95)
Krzemień
bottom 6.7 (0.0-12.4) 9.0 (3.0-24.0) 6.8 (4.0-8.4) 0.002 (0.001-0.004) 7.42 (6.68-7.84)
surf. 12.7 (8.8 - 17.9) 8.5 (2.8 - 15.5) 2.8 (0.5 - 13.2) 0.003 (0.000-0.014) 8.08 (7.30 - 8.55)
Miedwie
bottom 7.0 (0.8 - 12.9) 5.9 (2.0 -11.5) 9.4 (1.8 - 29.0) 0.000 (0.000-0.003) 7.46 (6.99 - 7.84)
surf. 11.1 (8.2-14.0) 15.4 (6.0-32.0) 8.1 (0.0-11.0) 0.006 (0.000-0.018) 8.09 (7.49-8.70)
Morzycko
bottom 3.3 (0.0-7.0) 9.2 (4.0-18.5) 6.8 (4.0-8.4) 0.001 (0.000-0.007) 7.50 (7.12-8.32)
surf. 11.1 93.9-14.00 6.6 (4.0-24.0) 7.5 (0.0-11.1) 0.016 (0.001-0.071) 8.12 (7.62-8.73)
Narost
bottom 3.4 (0.0-10.0) 13.6 (4.0-17.0) 9.4 (5.2-12.1) 0.002 (0.000-0.005) 7.48 (6.51-7.84)
surf. 9.1(1.9-13.6) 22.3 (8.0-42.5) 7.5 (0.0-11.6) 0.013 (0.003-0.076) 8.17 (6.90-9.42)
Wądół
bottom 3.4 (0.0-10.4) 11.5 (4.0-28.0) 11.7 (10.4-14.2) 0.005 (0.000-0.037) 7.59 (6.68-8.57)
surf. 11.3 (8.9-12.8) 21.2 (8.0-38.0) 7.1 (0.0-24.5) 0.004 (0.000-0.026) 7.92 (6.93-9.03)
Wisala
bottom 4.3 (0.0-12.0) 10.0 (5.0-23.0) 13.9 (2.8-23.8) 0.002 (0.000-0.008) 7.34 (6.79-7.83)
surf. 11.1 (8.2-13.2) 17.8 (2.0-54.0) 6.1 (0.0-10.5) 0.005 (0.001-0.039) 8.00 (7.58-8.91)
Woświn
bottom 6.3 (0.0-13.2) 8.9 (3.0-21.0) 7.7 (6.6-9.0) 0.005 (0.000-0.023) 7.66 (6.73-8.10)
Units: pH reaction without unit, the other indices in mg·dm-3.
185
Average values given first, maximum and minimum values in brackets

It was also found to be most variable in benthic waters, where it ranged from 2.0 (Ińsko Duże
and Miedwie Lakes) to 42.9 mg.dm-3 (Będzin Lake). In summer, when surface waters contained
the smallest amounts of dissolved carbon dioxide, its highest concentrations were determined
near the lake bottom (Tab. 3).
The content of suspension in the analyzed lakes was always higher in surface waters (from
4.0 to 54.0 mg.dm-3) than in benthic ones (from 2.0 to 28.0 mg.dm-3). The highest quantities
of suspension, frequently above 30 mg.dm-3, were recorded in summer in the surface waters
of the lakes Wądół, Wisala and Będzin. In the same season, the surface waters of the other
lakes were found to contain between 12 and 16 mg.dm-3 of suspension. In benthic waters,
in summer, the amount of suspension was 5-10 mg.dm-3, only on three occasions, in Będzin
and Wądół Lakes, exceeding 25 mg.dm-3. In winter, the concentration of suspension in benthic
waters ranged from 3 to 6 mg.dm-3. never going above 10.0 mg.dm-3. In surface waters, it was
typically 2-3 mg.dm-3 higher. As a rule, the surface waters in Miedwie, Ińsko Duże, Chłop Duży
and Jelenin were less abundant in suspension than in Będzin, Wądół, Ińsko Małe and Narost
Lakes (Tab. 3).
The oxygen conditions in the analyzed lakes are discussed below. In summer, in hyper-
trophic lakes Będzin, Wądół or in eutrophic Wisala Lake, the concentration of oxygen equal 4.0
mg.dm-3 was already recorded in the metalimnion, and in the hypolimnion it declined to trace
amounts while the benthic waters were completely deoxygenized. Oxygenation below 20%, at
which less tolerant fish find it difficult to breath, appeared in these lakes during the summer
stagnation in water layers as shallow as 6-8 m. Such oxygen conditions, unfavourable to fish,
were recorded in the analyzed lakes from early June to early November. In winter, the benthic
waters of Będzin and Wądół Lakes, and especially the latter one, had unfavourable oxygen
conditions, but in autumn and spring, they turned into suitable ones for fish.
In mesotrophic lakes Jelenin, Krzemień, Morzycko and Woświn, as well as in eutrophic
lakes Chłop Mały, Ińsko Małe, Wisala and Narost, oxygen in benthic waters occurred periodi-
cally. When the conditions were unfavourable, complete deoxygenation of benthic waters oc-
curred and the waters of the hypolimnion were considerably depleted of oxygen. In summer,
unfavourable oxygen conditions (such as oxygenation of waters less than 20% O2 saturation)
appeared in mesotrophic lakes at a depth of 10-12 m and in eutrophic lakes - in shallower
waters. The concentration of oxygen equal 4 mg.dm-3 appeared at depths of 6-8 and 4-6 m,
respectively. In benthic waters of the analyzed lakes, oxygen conditions which were unfavour-
able to fish persisted from the end of June to early October. In winter, it was only in Narost
and Wisala Lakes that the concentration of oxygen decreased to trace amounts or complete
deoxygenation occurred in the waters layers lying above the bottom.
In the recent years, Ińsko Duże and Miedwie Lakes have been characterized by oxygen
conditions suitable for fish which require much oxygen (Coregoninae). In the benthic waters of
these lakes, in the summer, autumn and winter, oxygenation only sporadically fell below 50%.
The amount of dissolved oxygen less than 4.0 mg.dm-3 (3.2 mg O2 .dm-3) occurred only once in
the deepest water layers of Ińsko Duże (in 1978). In turn, the benthic waters in Miedwie Lake
were completely deoxygenized in the 1980s and 1990s.
Chłop Duży Lake was characterized by the constant presence of oxygen in the benthic wa-
ters, but its amount was much smaller than in Ińsko Duże Lake. In the water at a depth of 12-14
186
m, the amount of dissolved oxygen fell below 4 mg.dm-3. Oxygenation equal 20% was typically
found below 18m, and 50% - in the thermocline 10-12 m deep (Tab. 3).
Discussion
The current state of lake eco-geosystems is a consequence of the process of eutrophi-
cation, which produces a significant, indirect effect on ichthyofauna by changing the
environmental conditions (Hartmann 1977, Prejs 1978, Zdanowski 1993, Opuszyński
1983, 1997, Leopold et al. 1986). Such changes include depressed water transparency,
disappearance of submerged plants, qualitative and quantitative changes in zooplank-
ton and benthic fauna, algal blooms, increased amounts of bottom sediments, worse
oxygen conditions especially in the deepest water layers as well as changes in the hy-
drochemical conditions in lakes, such as drastic changes in water pH (Wetzel 2002, Kajak
1979, Zdanowski 1993, Kalff 2002). When eutrophication begins, a moderate increase in
biological production occurs, which is a favourable development as it means increased
fish production. Later, however, many undesirable consequences of this process are
observed, such as mass development of phytoplankton organisms causing blooms of
mainly blue-green algae in surface waters, which reduce water transparency. Mass ac-
cumulation of these algae, excreting toxins, causes death of other organisms. Algal
blooms lead to the retreat to submerged plants due to deteriorating light conditions in
the littoral. Worse light conditions mean diminished abundance of benthic fauna, which
are food for fish that forage on these animals. In water bodies of high trophy, oxygen
conditions are disturbed; oxygen resources in the hypolimnion, and especially in the
profundal zone and benthic waters, are depleted, which leads to the disappearance of
deep-water fauna. Under anaerobic conditions, some chemical processes occur (ammo-
nification, denitrification), as a result of which such chemical substances as methane or
sulphur hydrogen are produced (Kajak 1979).
Modifications in fish assemblages are a sensitive indicator of changes in the trophy
of lakes (Opuszyński 1983, 1997, Zdanowski 1993). Due to eutrophication, some un-
desirable changes appear in the structure of ichthyofauna in particular water bodies
– commercially valuable species are supplanted by less desirable ones. In oligotrophic
lakes, ichthyofauna is initially dominated by species of the family Salmonidae, e.g. Euro-
pean whitefish Coregonus lavaretus (L.), vendace Coregonus albula (L.), lake trout Salmo
trutta m. lacustris (L.), which – as eutrophication progresses – are replaced by species of
the family Cyprinidae. This is a highly varied family, rich in species of different tolerance
to environmental and biocenotic conditions. The period of time when cyprinid fish domi-
nate lakes is the longest, and some significant changes appear then in the fish species
composition. Larger species, less numerous, which are important as food and game fish,
are supplanted by less attractive ones. As the ichthyofauna assemblages undergo such
qualitative changes, caused by increased trophy of lake waters, changes in the fishery
production volume follow; at first, more fish are captured, but then the increase in fish-
ery production is halted and in heavily eutrophic lakes, it becomes considerably smaller
(Prejs 1978, Opuszyński 1983, 1997, Zdanowski 1996).
In Poland, analyses of qualitative and quantitative changes in ichthyofauna triggered
by progressing eutrophication of lakes have been carried out, for example, by Iwaszk-
187
iewicz (1976) and Leopold et al. (1986). Most of our mesotrophic lakes have already
neared or reached the level of eutrophication which, when exceeded, will halt an in-
crease in the total fish biomass (Prejs 1978). Changes in the structure of ichthyofauna
caused by increased trophy of lakes have led to a situation when 65% of the lake fish
catches in West Pomerania today consist of cyprinids (mainly bream and roach) (Cabaj
2010).
The abiotic factors present in surface waters frequently produce highly varied effects
on fish, closely connected with the fish species’ sensitivity (Łukjanienko 1974). Fish are
highly adaptable, especially after staying for a longer time in an environment in which
harmful conditions appear in a concentration that does not cause pathogenic symp-
toms. The physiological and functional states of fish organisms, originally disturbed,
after some time will return to the normal ones.
Oxygen conditions are an important factor which shapes the habitat conditions of
fish. Many environmental factors affect the demand of fish for oxygen, e.g. tempera-
ture, pH, water dissolved O2 and CO2 as well as the rate of changes in their concentra-
tions. Previous adaptation of fish to such changes is equally important as much as the
tolerance of fish to unfavourable oxygen conditions (Kryteria… 1971, Opuszyński 1983).
Fish are more tolerant to a low concentration of oxygen at a pH close to neutral. For res-
piration processes, the best range of pH is from 6.0 to 8.5. as the water reaction changes
into alkaline or acidic, this tolerance decreases (Kryteria… 1971, Prost 1994).
Under normal conditions, the equilibrium between gas pressures in the fish’s blood
and in water is maintained. The difference between the external and internal pressure of
oxygen in trout should be about 20 mm Hg (Antychowicz 2007). The critical oxygena-
tion of water in a water body, necessary for keeping normal concentration of O2 in the
arterial blood of fish is different for different fish species. At the temperatures of 10, 15
and 20oC, it is, respectively, 8.0, 8.5 and 12.5% for roach Rutilus rutilus (L.), 14.6, 18.6
and 24.0% for bream Abramis brama (L.), 15.4, 25.0 and 30.5% for perch Perca fluviatilis
L., 19.4, 20.5 and 21.5% for pike and 26.0, 32.0 and 36.7% for rainbow trout (Guziur
1991). Even a small decline below the required minimum can reduce the presence of
coregonids and other salmonids (Szczerbowski 1993).
Oxygen demand of ichthyofauna alter depending on the species, gender, maturity
stage of gonads as well as the age and size of individual fish (Kryteria… 1971, Opuszyński
1983). The optimum oxygen concentration is 5-6 mg.dm-3, and when it falls below 5
mg.dm-3, foraging and body gain of fish decrease (Antychowicz and Wejman 1994). The
threshold, permissible concentration of dissolved oxygen, which enable a variety of fish
fauna to exist, is the value of 4.0 mg.dm-3 (Kryteria… 1971, Alabaster and Lloyd 1980).
Fish tolerance to depressed concentration of oxygen is short-lived and specific for
particular species and size groups; pike experiences breathing problems at 2-3 and dies
at 0.30-0.65 mg O2.dm-3; roach - at 2-3 and 0.7 mg.dm-3, perch – at 2.0-3.0 and 0.2-0.6
mg.dm-3; pikeperch – at 1.5-2.0 and 0.5-0.8 mg.dm-3, and bream – at 2.0-2.5 and 0.4-0.5
mg.dm-3 (Opuszyński 1983).
The highest oxygen consumption appears during reproduction – usually in both gen-
ders, although there could be a shift in time. Frequently, albeit not always, males use
188
more oxygen than females (Opuszyński 1983). In the water bodies where fish spawn,
the concentration of oxygen should be constantly at least 7 mg.dm-3. As the embryo in
an egg develops, and then during the early stage of development of a larva, oxygen
demand grows. Depressed concentration of oxygen in water, below the saturation level,
retards the embryonic development of cyprinid fish and delays their hatching (Szcz-
erbowski 1993),. As embryos develop, their sensitivity to oxygen deficits increases; it is
the highest prior to the hatching of larvae (Kryteria… 1971). The embryonic develop-
ment of some fish species can occur at an oxygen concentration below 2 mg.dm-3, but
such species as pike, bream or certa Vimba vimba (L.) need more than 4-5 mg O2 .dm-3
(Szczerbowski 1993). When the oxygen concentration falls to about 5-6 mg O2 .dm-3, the
larvae and fish fry of samonid fish can be retarded because their demand for oxygen is
much larger than that of older fish. Young fish are less tolerant to a low concentration
of oxygen but they are exceptionally able to gradually adapt to unfavourable oxygen
conditions (Antychowicz and Wejman 1994, Antychowicz 2007). Oxygen deficit in water
also depresses the fish’s ability to swim at a constant speed, especially the maximum one
(Kryteria… 1971, Alabaster and Lloyd 1980). The effects of hypoxia in ichthyofauna can
become particularly dangerous when the oxygen concentration in water drops rapidly.
A rapid decline in the concentration of oxygen down to the values tolerable under differ-
ent conditions, if accompanied by a rapid increase in the concentration of CO2, even to
some moderate values, can cause mass death of fish (Antychowicz and Wejman 1994).
It can be presumed that oxygen conditions in the analyzed lakes was a factor which
limited the occurrence of ichthyofauna. The threshold allowable amount of dissolved
oxygen in water at which varied ichthyofauna can exist is said to be rather low, no less
than 4 mg O2 .dm-3 (Kryteria… 1971, Alabaster and Lloyd 1980, Welch and Jacoby 2004).
Oxygenation of fish species which have high oxygen demand should be at least 70%
for Baltic whitefish, over 40% for pike-perch, over 15% for bream and slightly lower for
carp, eel and pike (Puczkow 1962)
In most of the examined lakes, fish which demand much oxygen and dwell in ben-
thic waters during their life cycle encountered unfavourable habitat conditions, includ-
ing poor foraging grounds (Hartmann 1977, Zdanowski 1993). A large decrease in the
content of oxygen in the meta- and hypolimnion forced benthophagous and plank-
tonophagous fish to forage in the epilimnion and littoral. This was also an undesirable
development for species which during their life cycle dwell in cool benthic waters, e.g.
Baltic whitefish. The most intensive body gain in fish occurs in summer; if the fish in the
examined lakes during that season could not feed on foraging grounds typical of their
species then their rate of growth was adversely affected and, consequently, the fishery
efficiency of a given lake was lower (Zdanowski 1993, Leopold et al. 1986, Welch and
Jacoby 2004). Thus, the unfavourable oxygen conditions could restrict possible forma-
tion of strong populations of fish species associated with the deeper parts of lakes. This
situation did not appear in lakes characterized by the best water quality, i.e. in Miedwie
and Ińsko Duże Lakes. In the recent years, they have belonged to mesotrophic lakes,
where the benthic and hypolminion waters offered good oxygen conditions, although
in the past these water layers had been found to lack oxygen.
189
The data presented in this paper demonstrate that oxygen conditions were the main
factor influencing the structure of fish assemblages in the analyzed lakes. This observa-
tion is verified by the opinion expressed in the literature on the predominant role of
oxygen in the formation of ichthyofauna in stagnant water bodies (Penczak 2000).
The respiratory functions of haemoglobin are significantly affected by the unit pres-
sure of carbon dioxide in fish blood. When this gas is released by fish, constant differ-
ence in the partial pressure of carbon dioxide must be maintained – the pressure of CO2
in blood should be higher than in the external environment. Any increase in the con-
centration of this gas in blood depresses the capacity of haemoglobin to bind oxygen.
Concentration of carbon dioxide in blood depends on the content of this gas in water
(Opuszyński 1983). If the concentration of carbon dioxide in water exceeds a certain
limit, metabolism in fish decreases and so does their demand for oxygen (Antychowicz
and Wejman 1994).
The sensitivity of fish to excess CO2 concentration in water depends on a fish species,
age, intensity of metabolism, current concentration of oxygen in water and alkalinity
of water. Fish of the highest oxygen demand, e.g. trout, are also characterized by the
highest CO2 production and the highest sensitivity to increased concentration of this
gas in water. Fish with higher metabolism (usually young fish) are more sensitive to
increased CO2 concentration in the environment. When their metabolism is low and the
water oxygenation high, salmonid fish can stand up to 70 and carp up to 300 mg CO2.
dm-3 in water without any visible adverse signs (Antychowicz and Wejman 1994). The
amounts of free CO2 found in the waters of the analyzed lakes did not pose a threat to
the ichthyofauna.
The amount of suspension in lake waters shapes the habitat conditions of aquatic
organisms which dwell in these lakes (Kryteria… 1971). Suspension can have an indirect
effect on fish, causing their mass death, retarding their growth rate, depressing the
tolerance to diseases or producing an unfavourable influence on the proper develop-
ment of spawn and larvae. It can also affect negatively the mobility and migrations of
fish, reduce amounts of available food and make fish catching more difficult (Kryteria…
1971, Szczerbowski 1993, Prost 1994). However, we lack evidence to claim that the
concentration of suspension in water below 25 mg.dm-3 has any negative effect on fish.
In principle, it is possible to keep fisheries on a good or average level in waters contain-
ing 25-80 mg.dm-3 of suspension. There is a very small difference in fish mortality at the
concentration of suspension equal 100 mg.dm-3 compared to pure water. The growth
of fish is also good when water contains 200 mg.dm-3 of suspension if fish have enough
food (Kryteria… 1971). Thus, it should be concluded that the amounts of suspension
determined in the examined lakes did not limit the presence of ichthyofauna.
The amount of water dissolved ammonia determines the effectiveness of the excre-
tion of NH3 from the fish’s circulatory system through the gills (Schmidt-Nielsen 1992)
and its release is completely blocked at pH above 10.5 (Antychowicz and Wejman 1994).
Toxicity of this compound to fish depends on a number of factors – it is directly propor-
tional to changes in the water reaction and temperature but reversely proportional to
oxygenation, water hardness and alkalinity (Kryteria… 1971, Łukjanienko 1974, Alabas-
ter and Lloyd 1980, Prost 1994, Welch and Jacoby 2004). Higher water pH and tempera-
190
ture cause ionized ammonia NH4 to transform into a toxic gas form NH3. Increased water
hardness depresses (via calcium ions) the permeability of cellular membranes and makes
it more difficult for toxins to penetrate inside cells. A low content of oxygen causes
higher fish mortality at threshold levels of NH3. It is presumed that when oxygenation
declines to 50%, the longevity of fish is 2/3 shorter (Antychowicz and Wejman 1994).
Resistance of fish to ammonia depends on a fish species, age, fitness and health, and
it is generally in accord with their sensitivity to oxygen deficit. It has been found out
that fish remaining under the influence of sublethal concentrations of ammonia acquire
enhanced resistance to subsequent, very high levels of this compound. A slow increase
in the concentration of NH3/NH4 in aquatic environment favours adaptation of fish and
leads to an increased level of toxicity of this compound. Under natural conditions, fish
are able to avoid toxic levels of ammonia (Kryteria… 1971, Alabaster and Lloyd 1980).
In general, it is claimed that 2-7 mg NH3.dm-3 causes death of most fish; the content
of 1.2-3.0 mg NH3.dm-3 may be lethal to more sensitive fish (e.g. salmonids) (Łukjanienko
1974). Because there are many factors which shape the toxicity of ammonia to fish, it
is said that the concentration of within 0.2-2 mg NH3.dm-3 is lethal, whereas the harm-
less one is 0.025 mg NH3.dm-3 (Kryteria… 1971, Alabaster and Lloyd 1980, Antychowicz
2007). Considering the fact that the concentration of non-ionized ammonia in the wa-
ters of the examined lakes rarely exceeded 0.025 mg NH3.dm-3, it can be concluded that
it was not a limiting factor for the fish dwelling in those lakes.
Another criterion taken for the evaluation of habitat conditions in the examined
lakes was water reaction. Observations have confirmed that fish appear in water of the
reaction equal 4-10 pH (Szczerbowski 1993, Antychowicz 2007), but it is assumed that
an allowable range is from 5 to 9. A change in water reaction that is sudden, big or
occurring in leaps causes death of fish (Kryteria… 1971). The harmful effect of a low
or high water reaction on fish consists of denaturation of proteins in the epithelium of
the skin and gills, which considerably impairs the exchange of oxygen and other com-
ponents between fish’s blood and the environment. Besides, in more acidic water, fish
are more vulnerable to diseases and their growth rate deteriorates (Szczerbowski 1993,
Prost 1994). The reaction of water also has a significant effect on toxicity of some chemi-
cal compounds (Łukjanienko 1974, Opuszyński 1983, Welch and Jacoby 2004).
It was only in Wądół Lake that the upper threshold level of water reaction tolerance
was exceeded in the surface waters during summer. In the other lakes, no such observa-
tions were made. In all the lakes, the increase in pH to maximum values in summer was
slow, therefore fish had a chance to adapt to the occurring changes. The highest water
reaction appeared in the surface water layer, being distinctly lower in deeper water lay-
ers. Noteworthy is the fact that fluctuations in pH are particularly dangerous to fish at
high temperatures; if the water reaction exceeds 6.0-8.5 then, even a small decrease in
the concentration of oxygen can cause mass death of fish (Antychowicz 2007). Based
on our study, it can be claimed that the concentration of hydrogen ions in the analyzed
lakes was not a factor that limited the presence of ichthyofauna.
We cannot, however, omit the question of the harmful effect on fish habitat condi-
tions produced by algae, especially blue-green ones., whose mass occurrence is charac-
teristic for waters of high trophy. Cyanotoxins produced by algae damage the liver and
191
nervous system of higher animals, therefore it can be assumed that they affect fish in
a similar fashion (Antychowicz 2007). This question, however, was outside the scope of
the present research, which dealt with abiotic factors.
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192
Habitat conditions for the
ichthyofauna in anthropogenic
Szmaragdowe Lake (Szczecin,
Poland) in 2009-2010
Arkadiusz Nędzarek, Agnieszka Tórz,
Jacek Kubiak, Sylwia Machula
Department of Hydrochemistry and Water Protection, Faculty of

Food Sciences and Fisheries, West Pomeranian University of Tech-
nology in Szczecin, ul. Kazimierza Królewicza 4,
71-550 Szczecin, e-mail: arkadiusz.nedzarek@zut.edu.pl
Abstract
The paper contains results of the hydrochemical analyses performed in Szmaragdowe
Lake in order to assess the habitat conditions for the lake’s ichthyofauna. The tests were
carried out in 2009-2010. Using standard methods, the temperature of the lake’s water, dis-
solved oxygen, concentration of biogenic elements, water reaction, total hardness, organic
matter, chlorophyll a, dry matter of seston and Secchi’s disc visibility were determined. It
has been demonstrated that the analyzed lake should be classified as a holomictic one with
two circulations (dimictic) and a bradymictic type in respect of the intensity of water mixing.
The oxygen conditions were typical of stratified eutrophic lakes. The high level of trophy has
been verified by an analysis made according to the method suggested by Carlson (1977),
which proved that the lake was on the borderline between mesotrophic and eutrophic lakes.
The hydrochemical parameters obtained for Szmaragdowe Lake indicate that it provides
habitat conditions for fish of intermediate to low environmental requirements.
Introduction
Due to man-made pressure, new water reservoirs have appeared in the terrestrial ecosys-
tem, either created for retention of water for municipal or commercial purposes or because
of unplanned and uncontrolled influx of water to excavation pits. Such reservoirs often un-
dergo the process of naturalization, and play an important role in shaping the landscape by
changing the morphology of an area, water relations, local climatic conditions, ecosystems
and habitats as well as the social and economic structure (Jaguś and Rzętała 2008).
In West Pomerania, an example of such a water reservoir, which appeared when water
flooded a former chalk pit, is Szmaragdowe Lake (Filipiak and Raczyński 2000). This reservoir
lies within the administrative borders of Szczecin, in Bukowa Forest Landscape Park (Fig. 1).
193
The aim of this study has

been to evaluate the hydro-
chemical conditions in Szma-
ragdowe Lake in the context
of the habitat conditions for
its ichthyofauna, and to de-
termine its trophic state us-
ing the method proposed by
Carlson (1977).
Methods Figure 1. Location of Szmaragdowe Lake and the site of water sampling
Szmaragdowe Lake was (S1) during the hydrochemical studies in 2009-2010.
created on 16th July 1925
when groundwater surged into a chalk pit. The lake’s water table has established at an altitude
of 42.2 m above sea level. The surface area of the reservoir is about 2.5 ha, its maximum length
is 275 m, the maximum width is 150 m and the maximum depth goes down to about 16 m.
The shores are formed as steep slopes, often resembling cliffs, and there is just a short section
to the south of the lake where the shore is flat. The reservoir’s bowl sinks deep into the marly and
chalky substratum. The land around the lake is about 30-40 m above the water table. The whole
area surrounding the lake is afforested. Szmaragdowe Lake is a water body with no tributary or
discharge streams and it has a small forest drainage basin. For a few years now, the water table
has been observed to decline. Szmaragdowe Lake is a near triangle in shape, and at its northern
tip there is a concrete bridge, which today is used as a vantage point.
Szmaragdowe Lake is not used for commercial fishing, but some unauthorized ob-
servations and information suggest that the presence of such fish species as the pump-
kinseed sunfish (Lepomis gibbosus), catfish (Silurus glanis), ide (Leuciscus idus), asp (As-
pius aspius) and rudd (Scardinius erythrophthalmus) has been recorded.
Field tests and laboratory examinations have been conducted in accord with the
methodology recommended by the APHA (1995). Samples of water were taken twice in
each season specific for the moderate climate. Water samples were collected from a site
situated in the middle of the lake about 15 m deep (Fig. 1) at the following depths: 0.5
m below the surface and then 3, 5, 8, 10 and 12 m as well as 0.5 above the bottom.
During the field tests, water temperature, water dissolved oxygen (DO) and water reac-
tion were measured using a multi-parameter probe HI 9828 HANNA Instrument. Secchi’s
discs were used to measure water transparency.
Titration methods were applied to assess the chemical oxygen demand with the di-
chromium method (CODCr) and total hardness in the sampled material. The dry matter of
seston was determined by the weight method. Total nitrogen was determined using a ni-
trogen analyzer manufactured by SHIMADZU TOC-VCSN. The other hydrochemical param-
eters were determined with colorimetric methods, using a UV-VIS spectrophotometer by
PerkinElmer, model LAMBDA 25, and the absorbance was measured at the recommended
wavelengths (λ). Ammonia nitrogen was determined with indophenol blue (λ=630 nm);
nitrite nitrogen was determined with sulfanilamide (λ=543 nm) and nitrate nitrogen was
determined as nitrites after initial reduction of nitrates to nitrites on a Cu/Cd column.
Inorganic nitrogen was calculated as a sum of ammonia, nitrite and nitrate nitrogen. Re-
active phosphorus was determined with the molybdenum method, using ascorbic acid as
a reductant (λ=882 nm); total phosphorus was determined as reactive phosphorus after
194
Habitat conditions for the ichthyofauna in anthropogenic Szmaragdowe Lake…
initial mineralization of a water sample with potassium persulphate in acid environment.

Chlorophyll a was determined after extraction with 90% acetone (λ =665 nm).
The results were submitted to statistical analysis. Pearson’s linear correlation coefficients
were calculated for correlations between the selected hydrochemical parameters, at a level of
significance α = 0.05, using the Statistica 9.0 software (StatSoft, Inc. 2009) for this purpose.
Results
During the studies, the temperature of water in Szmaragdowe Lake ranged within 0.8 to
19.9oC (Tab. 1, Fig. 2). The seasonal changeability of the water temperature was characterized
by the maximum values in the water layer down to 4 meters deep in the summer (from 19.9
to 17oC). Below, a large drop in temperature was recorded – of the gradient 2.2oC m-1 – the
thickness of this water layer was 5 m. Underneath, cool water of an average temperature of
5.1oC was lying. The lowest water temperatures were recorded in the wintertime, when the
lake was covered with permanent ice sheet from the end of December 2009 to mid-March
2010. During that time, the lowest water temperatures were measured in the top layer, directly
under the ice cover (about 0.8oC), whereas in the deeper water layers it was about 4oC. During
the spring and autumn circulation, complete homothermy was observed.
Table 1. Minimum, maximum, average (n=49) and standard deviation (SD) values
of the analyzed hydrochemical indices in the waters of Szmaragdowe Lake in 2009-2010.
Temper- Dry matter Chloro-

O2 CODCr
ature %O2 pH of seson phyll a
o
C mgO2 dm-3 mgO2 dm-3 mg dm-3 mg m-3
Minimum 0.8 0.0 0.0 6.54 24.6 16.6 0.91
Maximum 19.9 9.30 84.1 9.51 48.9 48.8 36.50
Average 6.8 3.55 30.08 7.88 37.1 27.9 9.14
SD 4.7 2.6 23.1 0.6 6.7 8.9 6.8
Reac- Total hard- Trans-

N-NH4+ N-NO3- Total N Total P
tive P ness parency
mgN dm-3 mgP dm-3 mval dm-3 m
Minimum 0.025 0.102 0.289 0.018 0.066 5.00 2.1
Maximum 1.168 0.789 2.639 0.508 0.586 7.00 4.5
Average 0.207 0.202 0.616 0.100 0.172 5.85 3.46*
SD 0.226 0.110 0.475 0.094 0.115 0.51 0.85
*n=7
The waters of the studied reservoir were characterized by a high changeability of its
oxygen conditions. The changeability scope of the dissolved oxygen and the percentage of
oxygenation spanned from the immeasurable values to 9.30 mgO2 dm-3 and 84.1%O2, and
the mean values for those indexes were respectively: 3.55 mgO2 dm-3 and 30.08 %O2 (Tab.
1). It was shown, at the same time, that through the majority of the study time, concentra-
tions of the oxygen dissolved in the water exceeded 4 mgO2 dm-3 in the surface layer with
the thickness of circa 8 m. In the deeper zones of the lake large oxygen deficits, with the
bottom zone deoxygenation, were noted (Fig. 3).
The organic matter, estimated with the chemical demand for oxygen (CODCr), was char-
acterised by a high level of concentration throughout the time of the studies and the mean
value of that index was 37.08 mgO2 dm-3 (SD=6.7) (Tab. 1). At the same time, a depth-de-
pendent increase of the CODCr values was noticed (Fig. 4). A statistical analysis has shown
195
that the chemical demand

was significantly co-re-
lated with the dissolved
oxygen (r=-0.49) and the
dry mass of the seston
(r=0.78). There was a pro-
portional dependency with
chlorophyll “a” but it was
not statistically significant
(r=0.22).
The concentrations
of chlorophyll “a” were
characterised by their high Figure 2. Seasonal water temperature (oC) changeability
changeability, from 0.91 to in Szmaragdowe Lake in 2009-2010.
36.50 mg m-3, with the mean
value 9.14 mg m-3 (Tab. 1).
Seasonal changeability was
characterised by maximum
values in the time of peak
vegetation and minimum
ones in winter (Fig. 5). A de-
crease in the concentration
of chlorophyll “a” with the
depth was noticed in the
vertical section. A statistical
analysis has shown a propor-
tional dependency of chloro- Figure 3. Seasonal water dissolved oxygen (CODCr, mg dm-3)
phyll “a” with the dissolved changeability in Szmaragdowe Lake in 2009-2010.
oxygen (r=0.21), and the correlations with temperature and water reaction were statistically
significant (r=0.69 and r=0.40 respectively). Index of the trophic state, calculated using the
Carlson method (1977) was TSIChl=64.
The water reaction it the studied lake spanned from 6.54 to 9.51, with the mean value of
7.88 (Tab. 1). Seasonal changeability of that index was characterised by its minimum value in
springtime and maximum value in summer (6.98 and 8.10 respectively). Through the major-
ity of the study time, the decrease of the water reaction with the depth was a constant trend
(Fig. 6). It was only in wintertime that the lowest pH values were noted in the surface area
(circa 7.0) and the maximum ones in the water layer between 5 and 8 m deep (circa 8.5).
A statistical analysis has shown a proportional correlation between the water reaction and
the water temperature, dissolved oxygen and chlorophyll “a” (Fig. 7) (r=0.44, r=0.50 and
r=0.40 respectively).
The transparency of water in Szmaragdowe Lake ranged from 2.1 to 4.5m, with an aver-
age equal 3.48 m (Tab. 1). It was demonstrated that this index was negatively correlated with
the concentration of chlorophyll a (r=-0.24). The index of the trophic state calculated accord-
ing to Carlson’s method (1977) was TSISD=45.
196
Concentrations of bio-
genic elements were high. For
instance, average concentra-
tions of inorganic nitrogen
and total nitrogen were 0.468
and 0.616 mg N dm-3, and
those of reactive and total
phosphorus were 0.100 and
0.172 mg P dm-3, respectively
(Tab. 1). In the nitrogen cycle,
it was observed that nitrate
nitrogen and ammonia nitro-
gen had equal shares in in-
organic nitrogen, but nitrate
nitrogen prevailed over am-
monia nitrogen in water lay- Figure 4. Seasonal chemical oxygen demand
in waters of Szmaragdowe Lake in 2009-2010.
ers containing more dissolved
oxygen. In water layers char-
acterized by oxygen deficits,
ammonia nitrogen prevailed
and its concentration tended
to increase in deeper waters
of the lake (Fig. 8). Statisti-
cally significant reversely pro-
portional dependences were
demonstrated of ammonia
nitrogen on dissolved oxygen
(r=-0.38) and ammonia nitro-
gen on water reaction (r=-
0.40) (Fig. 7).
The general tendency Figure 5. Dependence of chlorophyll a and water dissolved oxygen on the
for seasonal changes in water temperature in Szmaragdowe Lake in 2009-2010.
concentrations of inorganic
nitrogen and reactive phosphorus was characterized by decreasing concentration during
the peak vegetative season. The statistical analysis showed negative correlation between
reactive phosphorus and chlorophyll a (Fig. 9). The index of the trophic state according
to Carlson’s method with respect to total phosphorus concentration was TSITP=59.
The waters of Szmaragdowe Lake were characterized by high total hardness (on av-
erage 5.85 mval dm-3) (Tab. 1). In the subsequent seasons analyzed, slightly changeable
average values of this parameter were recorded, within 5.69 mval dm-3 (in autumn) to
6.33 mval dm-3 (in spring). In a vertical water column, water hardness tended to de-
crease with depth.
The content of dry matter of seston varied from 16.6 mg dm-3 to 48.8 mg dm-3 (Tab. 1).
The seasonal variability of this index was characterized by the maximum values in summer
197
Figure 6. Seasonal water reaction changeability in Figure 7. Correlations between water reaction,
Szmaragdowe Lake in 2009-2010. chlorophyll a and ammonia nitrogen in waters
of Szmaragdowe Lake (statistically
significant correlation coefficients)
and autumn (on average about 32 mg dm-3) and the minimum ones in winter and spring (on
average about 22 mg dm-3). In the vertical water column, an increase in the concentration of
seston’s dry matter was observed with increasing depth. This index was statistically signifi-
cantly correlated with chlorophyll a (r=0.49) and chemical oxygen demand (r=0.78).
Discussion
Water temperature is one of the major environmental factors which condition the
functions of aquatic ecosystems at all trophic levels. Water temperature depends on the
electromagnetic radiation which reaches water and on the factors which distribute the
absorbed radiation in the whole water volume (Wetzel 2001). Different distributions of
water temperature in lakes enable us to distinguish many types of lakes characterized
by particular temperature regimes (Skowron 2009). With respect to Szmaragdowe Lake,
the recorded thermal conditions suggest that it is a holomictic lake with two circulations
– spring and autumn ones (dimictic lakes) of the bradymictic type of the water mixing in-
tensity (Wiszniewski 1953, Hutchinson 1957, Kalff 2001, Wetzel 2001, Skowron 2009).
As the evaluation of the trophic state completed according the method proposed
by Carlson (1977) showed, Szmaragdowe Lake should be classified as a border one be-
tween mesotrophy (in respect of the visibility of Secchi’s disc) and eutrophy (regarding
Figure 8. Seasonal ammonia nitrogen (mg N dm-3) Figure 9. Dependence of chlorophyll a on inorganic
changeability in waters of Szmaragdowe Lake in nitrogen and reactive phosphorus in waters
2009-2010. of Szmaragdowe Lake in 2009-2010.
198
chlorophyll a and total phosphorus). Comparing the current evaluation of the trophic
state of this lake with the data from 1997-1998 (Raczyńska and Kubiak 2003), it should
be concluded that over the past ten years the trophic state has not changed. High fer-
tility of the water in this lake and thermal stratification conditioned the distribution of
water dissolved oxygen. During the summer stagnation, the oxygen curve had the shape
of a clinograde (complete deoxygenation of water was recorded at a depth of 8 m);
a clinograde oxygen curve is typical of stratified eutrophic lakes. In such lakes, photosyn-
thesis is the main source of oxygen in the euphotic zone and oxygen decomposition of
organic matter leads to complete depletion of water dissolved oxygen in deeper water
layers (Hutchinson 1957, Kalff 2001). In the analyzed lake, the occurrence of these proc-
esses is indicated by the recorded proportional dependence between DO and chlorophyll
a as well as the reversely proportional dependence between DO and CODCr.
Following Kudelska et al. (1994), it can be assumed that a negative feature of Szmarag-
dowe Lake is that it is not a discharge lake, which means that is accumulates substances
from the drainage basin. As a result, organic matter and biogenic substances accumulate
in the bottom sediments (which is confirmed by the recorded increase in the concentration
of organic matter and biogenic elements at deeper layers of the lake). In stratified lakes,
organic matter and biogenic elements accumulated in the intersticial layer above the bot-
tom and the sediments in the bottom undergo multi-directional biochemical transforma-
tions. However, due to the holomictic water mixing during the spring and autumn circula-
tion, they are not completely and permanently removed from the ecosystem and therefore
are a significant, internal source of biogenic elements (Zdanowski 1982, Sehgal and Welch
1991, Kalff 2001). This conclusion is confirmed by a very high primary production of the
lake, which can be assessed with an aid of chlorophyll a (Manuel and Zimmerman 1999,
Ostapenia et al. 2009). High primary production can be perceived as a self-limiting fac-
tor. As Kufel claims (2001), algae modify the light conditions in water, adversely affecting
water transparency. This may occur in Szmaradgowe Lake, where a reversely proportional
dependence was found between transparency and chlorophyll a. With the information
provided by Kufel and Kufel (1997) it can be concluded that algae in Szmaragdowe Lake
are an important component of seston, as a proportional relationship between chlorophyll
a and dry matter of seston was revealed.
The oxygen conditions recorded in Szmaragdowe Lake should be perceived as not
very suitable for ichthyofauna, especially for the fish species which have high environ-
mental requirements. It is assumed that the optimum oxygen concentration for fish is
5-6 mg O2 dm-3. When it falls below 5 mg O2 dm-3, foraging and body gains among fish
are limited (Alabaster and Lloyd 1982, Müller and Stadelmann 2004). The oxygen defi-
cit found in most of the water layers in the lake indicates that the living conditions for
benthophagous animals in that lake are very limited and suitable habitat conditions for
ichthyofauna appear in the epilimnion and littoral zones.
The negative influence of eutrophication on ichthyofauna is also reflected by a high
water reaction and high concentration of ammonia nitrogen (Alabaster and Lloyd 1982,
Müller and Stadelmann 2004). The waters in the analyzed lake were characterized by
quite a stable water reaction, with the average pH equal 7.88. The maximum values of
water reaction were recorded only during the peak vegetative season, which implies that
this parameter is affected by intensive photosynthesis. At the same time, the high total
hardness of water in Szmaragdowe Lake should buffer changes in pH (Kalff 2001). The
199
water reaction determined in our study should not affect the dissociation of ammonia
because when the pH value is below 8, the products of this reaction are ammonia ions,
which are harmless to fish (Rubin and Elmaraghy 1977, Meade and Watts 1995). As the
research demonstrated, when the water reaction increased, the concentration of ammo-
nia nitrogen in water decreased. The ranges of ammonia that are lethal to fish are quite
wide, for example rainbow trout tolerates about 0.2 mg dm-3 and perch can survive the
ammonia concentration as high as 1.2 mg dm-3 (Rubin and Elmaraghy 1977, Meade and
Watts 1995). Relatively high concentrations of ammonia nitrogen in Szmaragdowe Lake
were recorded in deeper layers of water, which were also characterized by the lowest pH
values. It can therefore be claimed that the recorded concentration of ammonia nitrogen
is at the threshold value of being harmful to ichthyofauna.
Summary
The analyzed lake shows an elevated trophic state, which has a negative influence on
the habitat conditions for zoocenoses, including ichthyofauna. The presence and size of fish
populations dwelling in Szmaragdowe Lake can be limited by the concentration of water
dissolved oxygen and content of ammonia. Thus, it is recommended to monitor the quality
of water in this lake and to work out a good method for its management and protection.
References
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Chesapeake Bay: Sources and reactivity. Estuaries 22(4): 980-994.
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Kufel I., Kufel L. 1997. Eutrophication processes in a shallow, macrophyte-dominated lake – nu-
trient loading to and flow through Lake Łuknajno (Poland). Hydrobiologia 324/343: 387-394.
Kufel L. 2001. Uncoupling of chlorophyll and nutrients in lakes – possible reasons, expected
consequences. Hydrobiologia 443: 59-67.
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crayfish, Cherax quadricarinatus. J. Shellfish Res. 14: 341–346.
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trophic lakes by oxygenation. Fisheries Management and Ecology 11, 251-260.
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trophic status. Freshwater Biology 54: 1312-1323.
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zecin Landscape Park. Acta Scientarum Polonorum, Piscaria 2(2): 97-116.
Rubin A.J., Elmaraghy G.A. 1977. Studies on the toxicity of ammonia, nitrate and their mixtures
to guppy fry. Water Research 11: 927–935.
Sehgal S. H., Welch E. B. 1991. A case of unusually high oxygen demand in a eutrophic lake.
Hydrobiology 209: 235-243.
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StatSoft, Inc. 2009. STATISTICA (data analysis software system), version 9.0. www.statsoft.com.
Wetzel R.G. 2001. Limnology. Lake and River Ecosystems, 3rd ed. Academic Press, San Diego.
Wiszniewski J. 1953. Uwagi w sprawie typologii jezior polskich. Polskie Archiwum Hydrobiologii 1(14): 11-23.
Zdanowski B. 1982. Variability of nitrogen and phosphorus contents and lake eutrophication.
Pol. Arch. Hydrobiol. 29(3-4): 541-597.
200
Preliminary studies on fishing
and transport of the sichel
(Pelecus cultratus L., 1758) for
reproduction under controlled
conditions
Roman Kujawa, Przemysław Pol*,
Andrzej Mamcarz, Grażyna Furgała-Selezniow
Department of Lake and River Fisheries, Faculty of Environmental

Protection and Fisheries, University of Warmia and Mazury in Olsz-
tyn , e-mail: reofish@uwm.edu.pl
* Institute of Animal Biology in Balice
Abstract
The sichel, for many years a species of a large coverage and very numerous in some
rivers, has become a threatened species, which now needs protection, not just fishing
restrictions but also fish fry stocking. Thus, it has become necessary to work out the bio-
technology for reproducing sichel and rearing up its larvae. The first stage was to catch
spawners from the natural environment. The fish were obtained in May and June from
the Vistula Lagoon, using stationary fishing gear (gillnets, traps). Much better results
were obtained using traps rather than gillnets. The captured spawners were transported
in polyethylene bags, in water with oxygen atmosphere and anesthetic, to the Aquarium
Laboratory of the Department of Lake and River Fisheries, at the UWM in Olsztyn, where
they were kept in specially adjusted tanks until spawning. All manipulations on fish must
be carried out very carefully and gently, given them an anesthetic (MS 222, 5 mg dm-3 of
water) so as to avoid the shedding of scales, which may lead to death of fish.
Introduction
The sichel (Pelecus cultratus L.) is the only representative of the fish native to Poland
whose body conformation resembles that of tropical flying fish. It has an extremely com-
pressed body with exceptionally large pectoral fins, set horizontally (Photo 1).
201
The territory over which

sichel once appeared used
to be large, covering the
systems of big rivers of the
basins of the Caspian Sea
(the Ural, the Volga), the
Black Sea (from the Kuban
to the Danube) and the Bal-
tic Sea (from the Odra to
the Neva Rivers) (Sabaneev
1959, Brylińska 2000). Un- Photo 1. Sichel (Photo R. Kujawa).
fortunately, in the recent years, it has been noticed that the coverage of this species is
drastically diminishing, especially in Europe (Witkowski and Heese 1996).
For many years, sichel was thought to be an undesirable species in inland fisheries,
and frequently perceived as the so-called “fish weed”. Back in the 18th c., Kluk (1780)
described it as fish for “common folk”, found in large numbers, especially in the Vistula
River. Nonetheless, there were regions in Poland (the Vistula Lagoon) where it had some
commercial importance (Benecke 1880, Filuk 1966). Recently, our opinion of this species
has changed diametrically, as evidenced by such actions as bans on its fishing imposed
in most coastal waters except the Vistula Lagoon. Studies on the state of ichthyofauna in
Polish rivers show that the abundance of sichel has diminished dramatically (Witkowski
1996, Witkowski et al. 2000).
According to the IUCN classification, sichel belongs to threatened species (the IUNC
code LC – least concern) (IUCN 2010). In the Polish Red Book of Animals, it can be found
among species threatened with extinction as a species of lower risk but near threatened
(code NT) (Głowaciński, ed. 2001). On the Polish Red List of lampreys and fish, as of
2009, sichel was listed as critically threatened (code CR) (Witkowski et al. 2009). Its
status since 1999, when it was considered an endangered species (code EN) (Witkowski
et al. 1999), has deteriorated. In Poland, it is under species protection except the waters
in the Vistula Lagoon (Journal of Law 2004). According to the literature data (Terlecki
2000), the Vistula Lagoon is still inhabited by a strong and quite stable population of
sichel, although its abundance constantly varies.
The world literature contains little information on the biology of sichel. Few publica-
tions deal mainly with the morphology and foraging behaviour of representatives of this
species in different water bodies (Berg 1949, Terlecki 1987, Chlopnikov 1992, Stolarski
1995, Krzykawski and Więcaszek 1997a,b). We lack information about its reproduction
under controlled conditions or rearing up juvenile stages. There are just single papers
describing the larval development of sichel (Koblickaja 1966).
Unless successful protection programmes, which will include possible mass rearing
of fish stock material under controlled conditions, are worked out, the status of sichel
can quickly change from “threatened” to “extinct”. Elaboration of the biotechnology for
its reproducing and rearing up of larvae is a prerequisite for maintaining and supporting
threatened populations of this fish. In order to achieve this aim, work on methods for
collecting sichel spawners from the natural environment has commenced.
202
Preliminary studies on fishing and transport of the sichel (Pelecus cultratus L., 1758)…
Obtaining spawners of sichel from the natural environment

Obtaining good quality spawners by catching them from the natural environment
is one of the major stages in controlled reproduction of sichel. Sichel spawners were
obtained in May and June 2010 from the Vistula Lagoon, by cooperating with profes-
sional fishermen. The time when spawners were caught (May and June) coincides with
the spawning season for this species in the wild. As in most fish species, the spawning
season of sichel is stretched in time, the reason being that while some individuals enter
reproduction others are not yet ready.
The fish were caught with stationary fishing gear: gillnets and traps (Photos 2, 3).
All manipulations were done very gently, because when removed from nets or traps,
spawners are at risk of losing scales, which may cause their death in a very short time.
Immediately after being
removed from the fishing
gear, the fish were placed
in containers with inten-
sive oxygenation (Photo 4).
They were also anesthetized
by adding to the containers
5 mg MS 222 per 1 dm3
of water. After reaching
the shore, the fish under-
went preliminary check-up. Photo 2. Trap set in the Vistula Lagoon (Photo R. Kujawa).
When females and males in
spawning condition were
found (i.e., with free flow-
ing eggs and milt, respec-
tively), eggs were obtained
by stripping and fertilized.
Than the fish were released
to the water body. Like-
wise, all the subsequent
manipulations with spawn-
ers were performed having Photo 3. Removing fish from traps (Photo R. Kujawa).
anesthetized them with MS
222 (5mg dm-3 of water)
(Photo 5).
Fertilized eggs were
transported to the Aquar-
ium Laboratory of the De-
partment of Lake and River
Fisheries at the University of
Warmia and Mazury in Ol-
sztyn. In order to transport
spawn in good condition, Photo 4. Fish being transported to the harbour (Photo R. Kujawa).
203
it needs to have constant

thermal conditions. This
can be achieved using port-
able fridges, in which tem-
perature can be maintained
at a set level. Having been
transported to a hatchery,
the fish eggs were incubat-
ed in Weiss jars.
The remaining fish,
Photo 5. Sichel in a solution with an anesthetic (Photo R. Kujawa).
which were not yet ready to
spawn, were moved into polyethylene bags, with water, tranquilizers (MS 222, 5mg dm-3
of water) and oxygen atmosphere and transported, like fertilized spawn, to the Aquarium
Laboratory of the Department of Lake and River Fisheries UWM in Olsztyn. The total time
(manipulations and transport) elapsing when fish were in bags with oxygen and anesthet-
ics was about 3 hours. Having been delivered to the Aquarium Laboratory, the spawners
were kept in spawning tanks (Kujawa et al. 1999), slightly modified to fit the require-
ments of sichel. The temperature of the water in the tanks was adjusted to the one in the
Vistula Lagoon during the fish catches, i.e. 14-16 oC. It was also very important to han-
dle the spawners very care-
fully and not to scare them
while keeping in the tanks
because they could hit the
walls of the tanks when
frightened and suffer from
a head injury (Photo 6).
The first attempts to ob-
tain sichel spawners were
not quite successful. Catch-
ing the fish into gillnets was Photo 6. A head damage of a sichel incurred
a gross misunderstanding, while the fish was kept in a tank (Photo R. Kujawa).
even though the nets were checked frequently. The fish died very quickly, many before
they were removed from the nets, while the ones which were still alive had lost many
scales. It should be mentioned here that scales of sichel are very delicately set. Thus, any
manipulation with these fish, without prior anesthesia, makes them shed scales, which
disturbs the mineral metabolism of the fish and leads to their death.
The fish caught into traps looked much better. But even in that case, when they are
removed from the traps, utmost care is recommended and holding them should be
reduced to the necessary minimum. Attempts to catch sichel specimens by hand often
ended up in the fish losing some of their scales. The first captured spawners of sichel
were not yet ready to spawn, which is why they were kept in the tanks.
In short, it can be concluded that obtaining sichel spawners ready to reproduce takes
much experience and time. All manipulations should be carried out very carefully and it
is best when the fish are anesthetized first.
204
Preliminary studies on fishing and transport of the sichel (Pelecus cultratus L., 1758)…
References
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stran. čast 2: 810-814.
Brylińska M. 2000. Ryby słodkowodne Polski. PWN, Warszawa.
Chlopnikov M. M. 1992. Pitaniye chiščnich ryb v Vislinskom Zalive Baltijskogo Morya. Voprosy
Ichtiologii 32: 171-176.
Filuk J. 1966. Szczątki ryb z grodziska kultury łużyckiej w Tolkmicku nad Zalewem Wiślanym.
Wiad. Archeolog. 32(1-2): 223-234.
Głowaciński Z. (ed.) 2001. Polska czerwona księga zwierząt – kręgowce. PWRiL, Warszawa.
IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. www.iucnredlist.org, accesed
on: 18.11.2010.
Journal of Law (Dziennik Ustaw) 2004. Nr 220 poz. 2237. Rozporządzenie Ministra Środowiska
z dnia 28 września 2004 r. w sprawie gatunków dziko występujących zwierząt objętych ochroną.
Załącznik nr 1. Gatunki dziko występujących zwierząt objętych ochroną ścisłą, z wyszczególnieniem
gatunków wymagających ochrony czynnej.
Kluk K. 1780. Zwierząt domowych i dzikich, osobliwie krajowych, historyi naturalney początki i
gospodarstwo. T. III. O gadzie i rybach. Warszawa.
Koblickaja A.F. 1966. Opredelitel molodi ryb delty Volgi. Izdatelstvo”Nauka” Moskva.
Krzykawski S., Więcaszek B. 1997a. Biometric characteristics of sabrefish Pelecus cultratus (L.),
saithe Pollachius virens (L.) and sea bass Dicentrarchus labrax (L.) from new localities in polish wa-
ters. Acta Ichtiol. Piscat. 2: 3-15.
Krzykawski S., Więcaszek B. 1997b. Nowe stanowisko ciosy Pelecus cultratus (Linnaeus, 1758)
(Pisces: Ciprinidae), bardzo rzadkiego gatunku w wodach północno-zachodniej Polski. New local-
ity of sabrefish Pelecus cultratus (L.1758) (Pisces: Cyprinidae), a very rare species in the northwest
Poland. Przegl. Zool. 41(1-2): 79-82.
Kujawa R., Kucharczyk D., Mamcarz A. 1999. A model system for keeping spawners of wild and
domestic fish before artificial spawning. Aquacult. Eng. 20: 85-89.
Sabaneev L.P. 1959. Żizn i łovlia presnovodnych ryb. Gosselchozizdat USSR, Kiev.
Stolarski J. 1995. Sichel (Pelecus cultratus L.) from the Vistula Lagoon. Bulletin of Sea Fisheries
Institute 2 (135): 11-21
Terlecki J. 1987. Wzrost ciosy Pelecus cultratus (L.) z Zalewu Wiślanego. Acta Acad. Agricult. AC
Techn. Olst. 15: 27-36.
Terlecki J. 2000. Poradniki ochrony siedlisk i gatunków Natura 2000 – podręcznik metodyczny.
Gatunki zwierząt (z wyjątkiem ptaków). Ciosa. Ministerstwo Środowiska, Warszawa: 234-236.
Witkowski A. 1996. Zmiany w ichtiofaunie polskich rzek: gatunki rodzime i introdukowane.
Zool. Pol. 41: 29-40.
Witkowski A., Błachuta J., Kotusz J. 1999. Czerwona lista słodkowodnej ichtiofauny Polski.
Chrońmy Przyr. Ojcz. 55(4): 5-19.
Witkowski A., Błachuta J., Kotusz J., Kusznierz J. 2000. Lampreys and fishes of the upper and
middle Odra basin (Silesia, SW Poland). The present situation. Acta Hydrobiol. 42: 283-303.
Witkowski A., Heese T. 1996. Ochrona rzadkich i zagrożonych ryb w Polsce, stan aktualny i
perspektywy. Zool. Pol. 41: 1-195.
ski: Czerwona lista minogów i ryb – stan 2009. The degree of threat to freshwater ichthyofauna of
Poland: Red list of fishes and lampreys – situation in 2009. Chrońmy Przyr. Ojcz. 65(1): 33-52.
205
Preliminary characterization
of milt of the salmon (Salmo
salar L.) from rivers in West
Pomerania
Katarzyna Dziewulska, Józef Domagała
Department of General Zoology, University of Szczecin,

ul. Felczaka 3c, 71-412 Szczecin,
e-mail: katarzyna.dziewulska@univ.szczecin.pl
Abstract
The purpose of this paper has been to evaluate the basic parameters of milt of the
Atlantic salmon (Salmo salar L.) obtained from 36 individuals captured during spawning
migration to the Wieprz and Rega Rivers. The analyzed parameters included: spermato-
zoa density, morphology, motility, parameters of spermatozoa motility, seminal plasma
osmotic pressure and pH.
The density of spermatozoa reached on average 16.5x109 mL-1 within the range of
12.8-26.1x109mL-1. The morphology of spermatozoa was good. Heads of spermatozoa
measured (length x width) 3.2±0.04 x 2.6±0.07μm and the average length of flagella
was 35.1±0.88μm. The percentage of motile spermatozoa in most individuals was over
70%. The spermatozoa of the salmon individuals in which their motility was over 80%
were characterized by very rapid movement. Their velocity 7s after activation was on
average 201.9±25.0μm s-1 (VCL) at the average linearity of 75.0±15.0%. The duration of
movement was on average 30.5±4.4s. The value of osmotic pressure of seminal plasma
in salmon reached on average 285.0±18.8 mOsm kg-1 and the pH was 7.9±0.1.
According to the measured parameters of semen, the quality of semen in most salm-
on migrating to coastal rivers in Western Pomerania was good.
Introduction
Once the salmon indigenous to Poland disappeared (Chełkowski 1986, Bartel 2002),
a Latvian herd from the Daugava River (the Dźwina Zachodnia) was used for its resti-
tution. At present, adult salmon fish return for spawning to stocked rivers in Poland.
The gametes obtained from such spawners are used for production of stocking mate-
rial (Bartel 2001). In fish breeding farms, the offspring is reared up to the smolt stage
207
because the restitution programme is mainly based on stocking the mouths of coastal
rivers with smolt fish (Bartel 1997, 2001). Sometimes younger fish are released to rivers,
such as alevin or summer fry (Domagała and Bartel 1995, 1997, Bartel and Domagała
1997, Domagała 2000, 2007). Maintaining a population of salmon in Polish waters relies
on conducting controlled reproduction and continuous stocking of watercourses (Bartel
2002, Witkowski et al. 2009). In turn, the abundance of obtained offspring largely de-
pends on the quality of gametes used for reproduction. It is therefore justifiable to per-
form evaluation and selection of gametes prior to making a decision on their fate. The
purpose of this study has been to evaluate basic parameters of semen of Atlantic salmon
from Latvia, which is acclimating to the environmental conditions in Poland.

Analyses were made on milt obtained from salmon spawners swimming to spawning
grounds in the Wieprza River (catches at the Polish Angling Association field station in
Darłowo) and the Rega River (catches at the Szczecin PAA field station in Trzebiatow). Se-
men was sampled from 33 males (21 individuals caught on 4.11.2004 and 12 individu-
als on 13.11. 2007) from the Wieprza and 3 individuals captured on 6 November 2007
in the Rega. The average body length of the examined salmon from the Wieprza River
was 76.1 cm within the range of 46-118 cm, and from the Rega – 76.4 cm (range 65-83
cm). Milt was sampled to plastic containers, having massaged the abdominal part of
live or stunned fish. Some material was obtained with a 5 cm catheter in order to avoid
contamination with faeces or urine. Samples were transported on a sheet of wet ice, at
the temperature of +2 to + 4oC, to the laboratory of the Chair of General Zoology, the
University of Szczecin, where they were submitted to analysis after 4 to 10 hours. Sper-
matozoa density was determined in a Bürker chamber, having diluted the material to
2 500x in 0.8% NaCl. Spermatocrit was assessed in hematocrit capillaries after centrifu-
gation (3 000x g) for 10 minutes. Motility was determined subjectively (visually) and ob-
jectively using the computer-assisted sperm analysis (CASA) system with triple replicates,
activating germ cells with Billard’s liquid supplemented with 0.1% BSA. Parameters of
the spermatozoa motility were established using a system of computer-assisted analysis
of motion of spermatozoa with the software (Sperm Class Analyzer) made by Microptic,
under a Nikon Eclipse 50i microscope and an objective lens with a negative contrast
phase 250x. Milt of the motility over 80% (n=10) as assessed with the subjective method
was submitted to the computer-assisted analysis. The movement of spermatozoa was
analyzed on the seventh second after activation with an aid of a recorded 0.5-second-
long film. The analyzed spermatozoa motility parameters were:
MOT – percentage of motile sperm cells
VCL – curvilinear velocity (μm s-1), (spermatozoa of VCL>20 μm s-1 were assumed to
be motile)
VSL – straight-line velocity (μm s-1)
VAP – average path velocity (μm s-1)
LIN – linearity (VSL/VCL x 100) (%)
STR – straightness (VSL/VAPx100) (%)
208
Preliminary characterization of milt of the salmon (Salmo salar L.)…
ALH – amplitude of lateral head displacement (μm)

BCF – beat cross frequency (Hz).
Duration of the spermatozoa movement was likewise measured.
The size of sperm cells was measured in a semen smear on microscope slide covered
with coverslip. Measurements of spematozoa heads were made for ten individuals (100
heads per indiv.) under a 100x lens; the length of a flagellum was measured for 30 in-
dividuals using a 40x lens and a Nikon Eclipse 80i microscope with the NIS-Elements BR
2.30 software package.
In seminal plasma (n=10), osmotic pressure was measured with a Knauer osmom-
eter, and the pH was determined with a Sentron 1001 pH-meter. Seminal plasma was
obtained by centrifuging milt twice at 10 000 x g for ten minutes at 4oC.

For restitution of salmon, fish stocking material of the same origin is released to the
Rega and Wieprza Rivers (Bartel and Domagała 1997, Bartel 2000, 2002), therefore, in the
further part of this paper, their milt is characterized jointly. In about 65% of individuals, the
recorded motility of spermatozoa was over 80%, with the average 71% within the range
0-99%. In few individuals, low spermatozoa density was recorded, below 6 x 109 mL-1, as
well as low osmotic pressure and motility below 10%. These individuals were excluded from
further analysis. In these males, milt was sampled without a catheter and it is likely that the
material was contaminated with urine, which depressed the density and osmolality of milt as
well as caused early activation of spermatozoa. In order to reduce the risk of urine contami-
nation of milt, whenever possible, it is recommended to sample it with a catheter.
Spermatozoa density in the examined salmon was close to the values reported by
other authors (Kazakov 1979, Aas et al. 1991). It reached on average 16.5±3.7 x 109
mL-1 (±SD) within the range 12.8 – 26.1 x 109 mL-1 (n=30). Spermatocrit of the analyzed
salmon was on average 36% within the range 28-50%. The morphology of spermatozoa
was good. The size of oval heads of sperm cells (length x width) was 3.2± 0.04 x 2.6 ±
0.07 μm, and the length of a flagellum was 35.0 ± 0.88 μm.
Spermatozoa of the salmon fish were characterized by rapid motility and their velocity
7 seconds after activation was on average 201.9±25.0 µm s-1 (VCL), while straightline ve-
locity VSL=148.8 ±22.9 µm s-1. These parameters are approximately the same as obtained
for rainbow trout (Oncorhynchus mykiss) by Tonimoto and Morisawa (1988) and for sea
trout (Salmo trutta m. trutta) (Dziewulska et al., unpublished data). In other salmonids,
spermatozoa move at a similarly high velocity. The mean curvilinear velocity (VCL) between
5 and 20 seconds after activation, was 110-115 µm s-1 for rainbow trout (Dietrich et al.
2007) and 140.1 µm s-1 for brown trout (Dietrich et al. 2007). Spermatozoa of acipenserids
reach a similar average velocity of 118-160 µm s-1 (Glogowski et al. 2004, 2008). In other
species of teleost fishes, spermatozoa attain lower velocities, e.g. carp (Cyprinus carpio)
97.1 µm s-1 (Dietrich et al. 2007), ide (Leuciscus idus) 63 µm s-1 (Kowalski et al. 2006a), dace
(Leuciscus leuciscus) 46 µm s-1 (Kowalski et al. 2006a). Spermatozoa of pikeperch (Sander
lucioperca) reach the velocity from 41.8 µm s-1 (Demska-Zakęś et al. 2005) to ca 90 µm s-1
(Kowalski et al. 2006a). Spermatozoa faster than those of salmon have been found only
209
in sturgeon (Acipenser fulvescens) (310-320 µm s-1, Toth et al. 1997) and turbot (Psetta
maxima) (230 µm s-1, Dreanno et al. 1998). Most spermatozoa of the analyzed salmon
moved with quasilinear trak at an average linearity (LIN) of 76.0±15.0% and straightness
(STR) of 82.0±13.9%. Spermatozoa of pikeperch, for example, are characterized by low
linearity (Kowalski et al. 2006b, Sarosiek et al. 2004, Demska-Zakęś et al. 2005). Amplitude
of lateral head displacement with respect to an approximate path (ALH) in the examined
salmon was on average 2.6±1.0 µm, and the frequency of crossing the curvilinear path
by the approximate path (BCF) was 7.8±2.9 Hz. For comparison, the flagellum beat fre-
quency at the onset of activation in rainbow trout was 55 Hz (Cosson et al. 1991).
Adenosine triphosphate (ATP) is the basic energy nucleotid for the movement of
a spermatozoa flagellum (Gatti et al. 1989). Concentration of ATP in salmon’s spermato-
zoa was on average 56.1±19.4 pmol 10-6 spermatozoa, and the total adenine nucleotide
content TAN was on average 94.8±22.5 pmol 10-6 while the value of adenylate energy
charge AEC=0.67±0.08 (Dziewulska et al. 2010). In salmonids, ATP available during
the motility phase is mainly derived from the accumulation of this nucleotide prior to
the movement phase because the efficiency of oxidative phosphorylation during motil-
ity is low (Christen et al. 1987). The ATP value given above is sufficient for an average
of 30.5±4.4 s of salmon spermatozoa making movement. In carp and African catfish,
a higher value of ATP is sufficient for a longer motility phase; in carp it lasts for 45-90 s
up to 2 minutes of movement (20oC) (Redondo-Muller et al. 1991, Perchec et al. 1995)
and in the latter species – for 40 s (28oC) (Mansour et al. 2003). In other species, ATP is
synthesized during the motility phase (Lahnsteiner et al. 1999). In Siberian sturgeon and
turbot, oxidative phosphorylation is more efficient, because following a rapid decline
in the concentration of ATP, this nucleotide remains on a constant level throughout the
second phase of motility (Billard et al. 1999, Dreanno et al. 1999). In Siberian sturgeon,
ATP is maintained on a constant level of about 40 pmol 10-6 spermatozoa during 2 min-
utes of movement (Billard et al. 1999). Spermatozoa of turbot accumulate the highest
amounts of ATP among examined fish species, which enables them to obtain the highest
initial velocity of spermatozoa recorded in fishes, i.e. 250 μm s-1. Moreover, during the
motility phase they renew ATP resources and the stable phase is maintained at a level of
about 67 pmol 10-6 spermatozoa for 10 minutes of movement (Dreanno et al. 1999).
The composition of seminal plasma has great influence on the quality of spermatozoa.
Seminal plasma osmotic pressure is a good indicator of semen being contaminated with
urea. Good semen should have osmotic pressure within 220-360 mOsm kg-1 (Lahnsteiner
et al. 1998), while lower values of this parameter may indicate contamination with urine
so such milt cannot be submitted to research or storage (Glogowski et al. 2000, Dietrich
et al. 2005). The value of osmotic pressure of seminal plasma from the analyzed salmon
was on average 285.0±307, and the average pH was 7.9±0.1 (7.8-8.1).
It should be kept in mind that selection of specimens for reproduction must be based
on good quality of gametes. In addition, the principle of maintaining biodiversity and
specific genetic characteristics of local populations, where fish create local herds, should
also be considered. Another issue raised is whether routine procedures of controlled
reproduction, taken for example competitiveness between spermatozoa, enable us to
preserve sufficiently the biodiversity of species (Ciereszko 2010, oral report).
210
Preliminary characterization of milt of the salmon (Salmo salar L.)…
Based on the examined parameters, it can be concluded that the quality of semen in
most of the analyzed salmon specimens was good, which ensures potential reproductive
capacity of salmon in Polish waters during natural as well as “artificial” spawning.
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212
Hormonal stimulation of
males and its effectiveness in
reproduction and propagation
of local populations of rheophilic
cyprinid fish
Beata Irena Cejko1, Radosław Kajetan
Kowalski1, Dariusz Kucharczyk2 ,
Daniel Żarski2, Katarzyna Targońska2,
Jan Glogowski1
1
Department of Gamete and Embryo Biology, Institute of Animal
Reproduction and Food Research of the Polish Academy of Sciences,
Olsztyn
2
Protection and Fisheries, University of Warmia and Mazury in Olsztyn
Abstract
The effect of hormonal treatments, i.e. LH-RH, Ovopel [(D-Ala6, Pro9-NEt)-mGnRH],
Ovaprim [(D-Arg6, Pro9-NEt)-sGnRH], carp pituary homogenate (CPH) and human chori-
onic gonadotrophin (hCG) on spermiation in males of rheophilic fish, such as asp (Aspius
aspius), barbel (Barbus barbus), nase (Chondrostoma nasus), ide (Leuciscus idus), dace
(Leuciscus leuciscus), chub (Leuciscus cephalus) and vimba (Vimba vimba) has been ana-
lyzed. The effectiveness of the treatments has been verified in terms of the total volume
of milt in millilitres (TVM), and volume of milt per kg of body weight of males (VOM),
concentration of spermatozoa in milt and their total number (TSP). Selected quantita-
tive and qualitative parameters of ide and barbel’s milt have also been determined in
dependence on the time after stimulation with Ovopel. Based on the reference data and
own studies, it has been demonstrated that hormonal stimulation has a positive effect
on the volume of milt, number of produced spermatozoa and their motility compared to
the control group. It has also been found out that males spawning in waters of higher
temperature, i.e. within 18-20oC (barbel) respond more quickly to hormonal stimula-
tion than the ones spawning in cooler waters, i.e. 12-14oC (ide). Economic and practical
213
considerations make Ovopel the most popular stimulant in the biotechology of fish re-
production, including rheophilic ones. According to some recent research, Ovaprim can
likewise be used to stimulate spermiation.
Introduction
Aquatic organisms are particularly exposed to unfavourable changes in the natural
environment, which are triggered by the progressing anthropogenic impact, that is the
effect man has on the surrounding environment. Due to such changes, the range of oc-
currence of many fish taxa has been limited, for example that of rheophilic fish, some of
which are at risk of becoming critically threatened species, e.g. barbel (Barbus barbus)
and ide (Leuciscus idus). Dace (Leuciscus leuciscus), asp (Aspius aspius) and chub (Leucis-
cus cephalus), in turn, are near threatened, but their abundance still remains on a proper
level. Nase (Chondrostoma nasus) and vimba (Vimba vimba) are close to becoming ex-
tinct in the near future (Kowalska and Zakęś 2010).
Biotechniques developed for fish reproduction are a powerful instruments used in
aquaculture in order to obtain mature gametes. The stock material bred under con-
trolled conditions is used for example to stock open waters, which ensures active pro-
tection of biodiversity and maintenance of the natural structure of ichthyofauna. Fish
stock releases are commercial in character but they also have ecological backgrounds.
Among numerous substances used to stimulate spermation of rheophilic fish, popu-
lar are natural gonadotropin-releasing hormones (Jamróz et al. 2008; Targońska et al.
2010), preparations containing analogues of gonadotrophin, i.e. Ovopel [(D-Ala6,Pro9-
NEt)-mGnRH+metoclopramide] and Ovaprim [(D-Arg6,Pro9-NEt) sGnRH+domperidone],
(Cejko et al. 2008; Krejszeff et al. 2008) as well as natural gonadotrophins, i.e. carp
pituary homogenate (CPH) or human chorionic gonadotrophin (hCG) (Kucharczyk et al.
1999, Krejszeff et al. 2008). The efficacy of each of the stimulants, expressed through
the volume of obtained milt and number of spermatozoa of satisfactory quality depends
on many factors, such as the type of a hormone administered to fish, its dose and the
time elapsing from hormonal stimulation to production of milt (Cejko et al. 2010). This
paper contains a synthesis of studies carried out for several years on the efficacy of hor-
monal preparations in stimulating spermation of rheophilic cyprinid fish which belong
to the Polish native ichthyofauna.
The abbreviations used in this paper stand for:
TVM – total volume of milt in millilitres (ml)
VOM – volume of milt in millilitres per kg of body weight of a male (ml kg-1 b.w.)
TSP – total number of spermatozoa in billions (x 109)
MOT – percentage of motile spermatozoa (%)
PRG – progressive sperm motility (%)
VCL – curvilinear velocity (μm s-1)
VSL – straight-line velocity (μm s-1)
ALH – amplitude of lateral head displacement (μm)
BCF – beat cross frequency (Hz).
214
Hormonal stimulation of males and its effectiveness in reproduction and propagation…
The genus Aspius

Asp
The genus Aspius in
Polish inland freshwaters
is represented by just one
species, i.e. asp (Fig. 1), list-
ed as a protected fish in the
Natura 2000 programme.
Figure 1 Males of asp selected for reproduction.
Being the only predatory
fish among cyprinids, it
plays an important role in
maintaining the trophic
pyramid in balance, which
is why it is considered as
one of the species with
a key function in bioceno-
sis. The biotechnique of
its reproduction has been
elaborated (Targońska et
Figure 2. Hormonal stimulation of asp males.
al. 2008, 2010), and until
now its spermiation has been stimulated mainly with synthetic preparations (Tab. 1).
After Ovaprim stimulation, the motility and concentration of spermatozoa in asp milt
reached higher values (62% and 10.65x109 ml-1, respectively) than when fish were
administered Ovopel (40% and 6.72x109 ml-1, respectively). The activity of acid phos-
phatase (AcP), lactic dehydrogenase (LDH) and β-N-acetyl-glucosaminidase (β-NAG)
was also higher after administration of Ovaprim (Cejko et al. 2008). The data pre-
sented above indicate that application of Ovaprim yielded better results although the
values obtained did not differ significantly from the ones obtained when Ovapel was
administered (P>0.05). It should be underlined that stimulation with either of these
stimulants causes spermiation in 100% of individuals (Żarski et al. 2008a), thus both
can be recommended to use in order to produce sperm of this species. Figure 2 illus-
trates hormonal stimulation of asp males.
Table 1. Selected hormonal agents, their doses and the time
of obtaining milt after hormonal stimulation of asp.
Hormonal Time after

Dose hormonal Author
agents stimulation
Ovopel 0.5 granule kg1 b.w. 48 h Cejko et al.2008

[(D-Ala , Pro9-NEt)-mGnRH]
6
1 granule kg b.w.
-1
36 h Żarski et al.2008a
Ovaprim 0.25 ml kg-1 b.w. 48 h Cejko et al.2008

[(D-Arg , Pro9-NEt)-sGnRH]
6
0.5 ml kg-1 b.w. 36 h Żarski et al.2008a
215
The genus Barbus

Barbel
Studies on the stimulation of spermiation in barbel have been carried out using
Ovopel (1 granule kg-1 b.w.), Ovaprim (0.5 ml kg-1 b.w.) and hCG (500 UI kg-1 b.w.).
The purpose has been to estimate the motility of spermatozoa, their concentration in
milt, total protein content
in seminal plasma and os-
molality of seminal plasma
depending on the adminis-
tered hormonal stimulant
(Cejko et al. 2009). Milt
was obtained 24 hours
after the stimulation (Fig.
3), and the control group
consisted of males which
received 0.9% of NaCl. The Figure 3. Collecting barbel milt.
values of all parameters in each experimental group ranged on a similar level, without
being statistically different (Tab. 2), and the positive influence of the hormonal prepara-
tions on spermiation of barbel was evidenced by the fact that no milt was obtained from
the control group.
Alavi et al. (2008) described changes in the quality of milt and sperm of barbel, pointing
to the fact that as the reproductive season progressed in time, both the volume of milt and
concentration of spermatozoa decreased from 0.4 ml and 18.8x109 ml-1 in March to 0.15
ml and 12.5x109 ml-1 in May. Likewise the ionic composition of sperm plasma undergoes
modifications, which leads to its altered osmolality and morphological parameters of sper-
matozoa. The observed decrease in the volume of milt and inferior quality of spermatozoa
are most probably caused by the ageing of spermatozoa once the spawning season is over.
Changes in the sperm of barbel are also observed depending on the time elapsing
from hormonal stimulation, an event which was confirmed by analyzing quantitative
and qualitative parameters of milt obtained 12, 36 and 60 h after stimulation with
Ovopel
(1 granule kg-1, Fig. 4).
The control group com-
prised fish administered
0.9% of NaCl. The highest
TVM and VOM were deter-
mined 12 h after stimula-
tion and they were signifi-
cantly higher (P<0.001)
than the volume found in
the control group (Tab. 3).
36 and 60 h after stimu-
lation, the volume of ob- Figure 4. Granules of Ovopel used for hormonal stimulation
of males of rheophilic fish.
tained milt decreased, but
216
the differences were not statistically significant compared to 12 h (P>0.05). The highest
concentration of spermatozoa was found in the milt collected from the control fish,
whereas in the fish stimulated with hormones, extension of the time lapse from 12 to 36
h led to hydration (liquefying) of milt and a significant decrease in the sperm concentra-
tion due to the effect produced by the hormones (Tab. 3). The highest TSP values were
recorded after 12 h, but when the tests were carried out 36 and 60 h after stimulation,
no significant increase in the number of obtained spermatozoa was observed (P>0.05)
(Tab. 3).
Characteristics the motility of barbel sperm depending on the time lapse after hor-
monal stimulation were analyzed with the CASA (Computer Assisted Sperm Analysis)
system and according to the following motility parameters: MOT, PRG, VCL, VSL, ALH
and BCF. No significant in-
crease (P>0.05) in the val-
ue of MOT in time was ob-
served after administration
of Ovopel, but the value of
PRG was noticed to have
risen significantly 60 h fol-
lowing stimulation versus
30 h afterwards (P<0.01)
(Figs 5a,b). The highest to-
tal velocity and straight-line
velocity were determined
after 12 h (228.4 μm s-1
VCL and 167.9 μm s-1 VSL),
and these results were sig-
nificantly different from
the ones found after 36 h
(P<0.01) (Figs. 5c,d). The
values of ALH (1.93 μm) as
well as BCF (11.46 Hz) were
also the highest after 12 h
and differed significantly
from the values determined
after 36 h in the case of ALH
(P<0.001) and after 60 H Figure 5. Percentage of motile spermatozoa (a), progressive motility of
spermatozoa (b), curvilinear velocity (c), straight-line velocity (d) ampli-
for BCF (P<0.001) (Fig. 5e,
tude of lateral head displacement (e) and beat cross frequency (f) in the
f). The CASA estimated pa- milt of barbel determined in the control group (injection of 0.9% NaCl)
rameters may suggest that and after 12 h, 36 h and 60 h following hormonal stimulation. Bars
marked with different letters are significantly different.
milt 12 h after Ovopel stim-
ulation should be selected for fertilization of barbel eggs. The parameters characterizing
motility of spermatozoa as well as the values of TVM, VOM and TSP indicate that this is
the most suitable time for this species.
217
Table 2. Values of the basic quality of barbel milt obtained during a reproductive season after
stimulation of spermiation with Ovopel, Ovaprim and hCG (Cejko et al. 2009).
Motility of Concentra- Osmotic Total pro-

Groups spermatozoa tion pressure tein
(%) (x109 ml-1) (mOsm kg-1) (mg ml-1)
Ovopel
68±35,8 13,2±2,5 273±14,7 1,4±0,5
[(D-Ala6, Pro9-NEt)-mGnRH]
Ovaprim
76±10,3 11,2±2,8 270±4,9 1,2±0,3
[(D-Arg6, Pro9-NEt)-sGnRH]
hCG 75±13,7 13,0±2,5 277±22,4 1,5±0,5
Table 3. Values of TVM, VOM, TSP and concentration of spermatozoa in milt after
stimulating spermiation of barbel with Ovopel (1 granule kg-1), * injection of 0.9% NaCl.
Time after hormonal TVM VOM Concentration TSP

stimulation (ml) (ml kg-1 m.c.) (x109 ml-1) (x109)
*
Control 1,67a 18,25a 14,73a 24,89a
12 h 3,10b 37,24b 12,37b 37,85b
36 h 2,34ab 25,90ab 12,36b 29,14ab
60 h 2,27 ab
26,43 ab
14,06ab
31,09ab
The genus Chondrostoma

Nase
Nase belongs to the species at risk of becoming extinct in the wild in near future,
and therefore it requires thoughtful actions and protective treatments for the sake of
preserving biodiversity, using methods and instruments that aquaculture provides us
with (Targońska et al. 2008, Kujawa et al. 2010). The world literature lacks detailed data
on the quality of nase milt (Lahnsteiner et al. 2000). Under controlled conditions, nase
males produce sperm without hormonal stimulation (Żarski et al. 2008b), but the volume
of milt obtained from young males is small (100-500 μl, Cejko, unpublished data). As
we are short of data regarding hormonal stimulation of nase males, it is difficult to con-
clude whether administration of hormones can significantly raise the volume of sperm
and the number of spermatozoa. Lahnstener et al. (2000) cited that the concentration
of spermatozoa in nase milt ranged from 10-20x109 ml-1. Our own studies suggest that
the values of this parameter in nase are somewhat lower, i.e. within 4.30-12.90x109 ml-1
(n=14), whereas the values of TSP are 2.07x109 (Cejko, unpublished data). Nase males
anesthetizia procedure is shown in Figures 6 and 7.
The genus Leuciscus
Ide
For the stimulation of spermiation of ide (Fig. 8), a variety of hormonal agents have
been used (Tab. 4), testing the effect of some doses on the volume of milt, number of
spermatozoa and their motility. After stimulation with CPH or hCG, motility parameters
did not change significantly from those obtained when fish was stimulated with Ovopel
(Kucharczyk et al. 1999). Jamróz et al. (2008) report that after stimulation with Ovaprim,
218
the motility of spermato-

zoa was significantly high-
er than the motility detect-
ed after administration of
Ovopel (P<0.05). A similar
relationship was discovered
by Kucharczyk et al. (2007),
who analyzed the effect
of Ovaprim on reproduc-
tion of wild and ornamen-
tal form of ide (Leuciscus Figure 6. Anesthetizing nase males with 2-phenoxyethanole.
idus Auber. Ofrus). When
LH-RH was used, the mo-
tility of spermatozoa was
significantly higher (81%)
then after stimulation with
Ovopel (67%) (P<0.05)
(Jamróz et al. 2008).
Analogously to barbel,
the volume of milt obtained
from ide (Fig. 9) and the
number of spermatozoa Figure 7. Preparing nase males to collecting milt.
depended on the time fol-
lowing stimulation, but due
to a lower temperature of
water typical of the repro-
duction of this fish species,
this time is considerably
longer. After administra-
tion of Ovopel in a standard
dose, i.e. one granule kg-1
body weight, the highest
values of TVM and VOM in Figure 8. Hormonal stimulation of ide males.
ide were found after 84 h.
Likewise, the values of TSP
peaked at that time (Tab. 5).
Keeping males longer than
84 h after hormonal stimula-
tion is impractical, both be-
cause there is no significant
increase in the volume of milt
or number of spermatozoa
Figure 9. Ide milt collected into a syringe.
219
afterwards and because of the need to heat up large amounts of water for a longer time
(Cejko et al. 2010).
Table 4. Selected hormonal agents and their doses used to stimulate spermiation of ide.
Hormonal agents Dose Author

natural gonadotropin-releasing hormones
LH-RH 20 μg kg1b.w. Jamróz et al.2008
synthetic analogues of gonadotropin-releasing hormones
Kucharczyk 2002; Kucharczyk et al.1999,
0.5 granule kg1b.w.
Ovopel 2000; Targońska-Dietrich et al.2004
[(D-Ala6, Pro9-NEt)-mGnRH] Kucharczyk et al.2007; Jamróz et
1 granule kg1b.w.
al.2008; Cejko et al.2010
Ovaprim
0.5 ml kg1b.w. Kucharczyk et al.2007; Jamróz et al.2008
gonadotrophins
CPH 2.0 mg kg1b.w. Kucharczyk 2002; Kucharczyk et al.1999
hCG 1000 UI Kucharczyk 2002; Kucharczyk et al.1999
Table 5. Values of TVM, VOM, TSP and concentration of spermatozoa in milt of ide after
stimulating spermiation with Ovopel (1 granule kg-1) (Cejko et al. 2010), * injection of 0.9% NaCl.
Time after hormonal TVM VOM Concentration TSP

stimulation (ml) (ml kg-1 b.w.) (x109 ml-1) (x109)
*
Control 0.17a 0.78a 8.75a 1.41a
36 h 0.35ab 1.52ab 9.05a 3.28ab
60 h 0.65b 2.44ab 11.86a 7.00b
84 h 0.70 b
3.03 b
11.29 a
7.03b
108 h 0.61 ab
2.78 b
11.04 a
6.48b
Dace
Under controlled conditions, it is possible to obtain milt from dace both during the
spawning season (Kucharczyk 2002, Targońska-Dietrich et al. 2003, Cejko, unpublished
data) and outside that time (Kucharczyk 2002, Kupren et al. 2003). Hormonal stimulants
and their doses, which have been used so far to induce spermation in fish of this species
are specified in Table 6. During the reproductive season, the motility of spermatozoa
assumed similar values after stimulation with CPH, Ovopel, Ovopel+CPH and in the con-
trol group (61-71%, P>0.05) (Targońska-Dietrich et al. 2003). Also, the values of VOM
in each group were shaped on a similar level, i.e. 0.6-0.7 ml kg-1 b.w. (P>0.05). Lower
values of VOM were found by Kupren et al. (2003) outside the ide’s reproductory sea-
son, when spermiation was induced using CPH or Ovopel. In both groups, VOM was 0.3
ml kg-1 b.w., whereas the motility of spermatozoa was 70%. These values did not differ
significantly from the ones determined for the control group (0.2 ml kg-1 b.w., P>0.05);
differences, however, were observed in the motility of spermatozoa between the stimu-
lated groups and the control one (52%, P<0.05).
220
In subsequent studies on ide males, the effect of Ovaprim and LH-RH was tested, and
the usefulness of these preparations was determined according to the volume of milt
and number of produced spermatozoa. Milt was obtained during the spawning season
48 h after stimulation and the results are presented in Table 7. When comparing the
effectiveness of the selected hormonal stimulants, it can be concluded that the highest
values of TVM, VOM and TSP were achieved after stimulation with Ovaprim, whereas the
lowest ones – after injections of hCG and LH-RH. Hypophysation of males led to hydra-
tion of milt comparably to the effect produced by Ovaprim stimulation, and this can be
confirmed by the values of sperm concentration ranging in both cases on a similar level,
not significantly different (P>0.05) (Tab. 7). Because of the higher values of TSP, TVM
and VOM achieved after administration of Ovaprim to males, this preparation should be
recommended for induction of spermiation in males of this species.
Table 6. Selected hormonal agents and their doses used to stimulate spermiation of dace.
LH-RH 100 μg kg1 b.w. Cejko, unpublished data

synthetic analogues of natural gonadotropin-releasing hormones
Kucharczyk 2002; Kupren et al. 2003;
0.5 granule kg1 b.w.
Ovopel Targońska-Dietrich et al. 2003
[(D-Ala6, Pro9-NEt)-mGnRH] Kucharczyk 2002; Cejko, unpublished
1 granule kg1 b.w.
data
Ovaprim
0.5 ml kg1 b.w. Cejko, unpublished data
gonadotrophins
2.0 mg Kucharczyk 2002; Kupren et al. 2003
CPH
3.0 mg Cejko, unpublished data
500 UI Cejko, unpublished data
hCG
1000 UI Kucharczyk 2002
Table 7. Values of TVM, VOM, TSP and concentration of spermatozoa in milt after stimulating spermiation
of dace with selected hormonal agents, * injection of 0.9% NaCl.
TVM VOM Concentration TSP

Hormonal agents
(ml) (ml kg-1 b.w.) (x109 ml-1) (x109)
*
Control 0.23a 3.01a 6.29d 1.41a
Ovopel 0.58ab 6.32b 5.35cd 3.08bc
Ovaprim 1.01 c
11.50 c
4.13 ab
4.07c
hCG 0.41ab 4.78ab 5.21bcd 2.06ab
CPH 0.71 bc
7.44b
3.25 a
2.30ab
LH-RH 0.45ab 5.12ab 4.91bc 2.15ab
221
Chub
In chub, spermiation was induced with an aid of the preparations used previously in
fish of the genus Leuciscus, and the applied doses are specified in Table 8. In an experi-
ment run by Krejszeff et al. (2008), the efficacy of CPH, hCG and Ovopel was determined
according to just two parameters, i.e. percentage of motile spermatozoa and values of
VOM. Motility was similar in each group, assuming the values 80-90% and not being sta-
tistically significantly different (P>0.05) from the values obtained in the control group
(60-70%). The highest values of VOM were determined after hypophysation of males
(3.7 ml kg-1 b.w.); lower VOM appeared in the control group (2.4 ml kg-1 b.w.). Differ-
ences between these values were statistically significant (Krejszeff et al. 2008).
Twenty-four hours after stimulation, the highest TVM values were detected in fish
stimulated with Ovaprim and in the group receiving CPH, whereas the lowest ones ap-
peared in the control group (P<0.01) (Tab. 9). In the group which was administered Ov-
aprim, the concentration of spermatozoa in milt was the lowest, which suggests that the
milt underwent hydration following the application of the stimulant. Motility of sperma-
tozoa was 60% after Ovopel, 52% after Ovaprim, 58% after CPH, 59% after LH-RH and
36% in the control group. Significant differences in values of sperm motility were found
between the control group and the LH-RH stimulated one (P<0.05). Hormonal stimula-
tion of chub males and milt collected to test tubes are illustrated in Figures. 10 and 11.
Table 8. Selected hormonal agents and their doses used to stimulate spermiation of chub.

LH-RH 50 μg kg1b.w. Cejko, unpublished data
synthetic analogues of natural gonadotropin-releasing hormones
Ovopel Krejszeff et al.2008, Cejko,
0,5 granule kg1b.w.
[(D-Ala6, Pro9-NEt)-mGnRH] unpublished data
Ovaprim
0.25 ml kg1b.w. Cejko, unpublished data
gonadotrophins
Krejszeff et al.2008, Cejko,
CHP 2.0 mg kg1b.w.
unpublished data
hCG 1000 UI Krejszeff et al.2008
Table 9. Values of TVM, VOM, TSP and concentration of spermatozoa in milt after stimulating
spermiation of chub with selected hormonal agents, * injection of 0.9% NaCl.
TVM VOM Concentration TSP

Hormonal agents
(ml) (ml kg-1 b.w.) (x109 ml-1) (x109)
*
Control 2.88ab 10.80ab 10.26b 28.83ab
Ovopel 4.51bc 15.23bc 8.19ab 36.38ab
Ovaprim 5.88 c
19.24 c
5.47 a
33.08ab
CPH 5.39c 19.61c 8.12ab 42.84b
LH-RH 2.46a
8.95 a
9.68b
23.57a
222
Figure 10. Hormonal stimulation of chub males. Figure 11. Chub milt collected into test tubes.
The genus Vimba

Vimba
This is the only cyprinid species which undertakes anadromous spawning migra-
tion, and in the wild faces a high risk of becoming extinct in near future. As in the case
of nase, there is very little information about the quality of its milt. Hliwa et al. (2003)
determined some qualitative and biochemical parameters of vimba sperm after stimula-
tion with Ovopel. Motility of spermatozoa was determined at a level of 50-80% and the
average time of spermatozoa being motile was 26s. The sperm concentration was about
15-17x109 ml-1.
Summary
Hormonal stimulation is an effective tool used in reproduction of many fish species,
including rheophilic cyprinids. An advantage of hormonal stimulants is the possiblity to
obtain milt outside a reproductive season, but other factors, such as adequate water tem-
perature, are necessary to achieve success (Kucharczyk 2002, Kupren et al. 2003). In rhe-
ophilic fish, the time from injection of hormones to obtaining milt is measured in hours,
and - depending on a species – the highest volume of milt (TVM, VOM) and number of
spermatozoa (TPS) are obtained after 12 h in barbel (Cejko, unpublished data) up to 84 h
in ide (Cejko et al. 2010). These results suggest that males spawning in warmer waters, i.e.
within 18-20oC (barbel) respond more quickly to hormonal stimulation than species which
spawn in cooler waters, i.e. 12-14oC (ide). It is interesting to notice that hormonal injec-
tions not always lead to a significant increase in TVM compared to a control group, a fact
that is most often observed in smaller fish species, i.e. dace. In domesticated fish, eggs and
milt can be obtained without hormonal stimulation, provided that the maintained photo-
thermal conditions are suitable for a given species (Krejszeff et al. 2009).
Gonadotrophins are less often used in reproduction of rheophilic fish, although this
method of hypophysation, applied since the 1930s, is still in use (Krejszeff et al. 2008).
Chorionic gonadotrophin, i.e. hCG, is not very effective either, especially in the fish of
the genus Leuciscus. At present, the main role in reproduction of cyprinids is played by
synthetic preparations, which contain analogues of gonadotropin-releasing hormones
in combination with dopamine inhibitors. The most popular in fish hatcheries are two
products, known under commercial names as Ovopel and Ovaprim. With respect to rhe-
223
ophilic cyprinid fish, stimulation with Ovopel causes a significant increase in the volume
of obtained milt and number of spermatozoa compared to the control, soon after the
treatment (Kucharczyk et al. 1999, Krejszeff et al. 2008, Cejko et al. 2010). Satisfactory
results are also obtained when males are administered Ovaprim; on many occasions, the
results are superior to those achieved when Ovopel is given (Cejko et al. 2008, Jamróz et
al. 2008). Due to a strong mechanism inhibiting the release of gonadotrphins from the
pituary, it seems that administration of preparations which do not contain antagonists
of dopamine receptors, i.e. metoclopramide or domperidone, is not very successful.
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225
Effect of cryoprotectants
on activity of some enzymes
and motility parameters of
spermatozoa of rainbow trout
(Oncorhynchus mykiss) and
Siberian sturgeon (Acipenser
baerii)
Beata Sarosiek, Radosław Kowalski
Department of Gamete and Embryo Biology, Institute of Animal

Reproduction and Food Research of the Polish Academy
of Sciences, ul. Tuwima 10, 10-747 Olsztyn, Poland,
e-mail: b.sarosiek@pan.olsztyn.pl
Abstract
This paper presents the effect of some cryoprotectants (DMA, DMSO, methanol and
glycerol) on the activity of enzymes in the seminal plasma and extracts from spermatozoa of
rainbow trout and Siberian sturgeon. The activity of acid phosphatase from rainbow trout’s
sperm extracts increased in the presence of DMA and glycerol. The same enzyme extracted
from spermatozoa of Siberian sturgeon did not change their activity in the presence of
any cryoprotectants. In turn, β-N-acetylglucosaminidase proved to be very sensitive to ad-
dition of DMA, DMSO and glycerol, which caused decrease in their activity. The activity of
arylsulfatase from sturgeon’s seminal plasma decreased in the presence of the cryoprotect-
ants, in contrast to that extracted from its spermatozoa. The analysis of the sperm motility
parameters with and without cryoprotectants enabled us to conclude that an addition of
methanol or glycerol of trout’s spermatozoa affected most of these parameters. In the case
of sturgeon’s spermatozoa, it was only glycerol that affected their motility parameters.
Introduction
In the world literature, papers on biochemical assays of fish semen are relatively
scarce compared to the information on mammals. Considering the fact that fish are the
227
largest and highly differentiated group of vertebrates, the knowledge of biochemical

aspects of fish reproduction needs to be expanded, all the more so as the subphylum of
the teleost fish (Teleostei) is highly differentiated not only in the way they spawn but
also in the structure of their gametes.
Spermatozoa of most animals contain a characteristic organelle, such as an acrosome,
found in the apex of the head of a spermatozoon. The function of the acrosome is to en-
able the spermatozoon to fuse with an oocyte and penetrate its outer layer. During fertil-
ization, an acrosomal reaction occurs, when enzymatic proteins present in the acrosome
are released/activated. Most of the enzymes present in the acrosome belong to the class of
hydrolases. The following can be named: acrosin, arylsulfatases (AS), alkaline phosphatase
and acid phosphatase (AcP), β-glucuronidase or β-N-acetylglucosaminidase (β-NAGase).
Fish which belong to the order Salmoniformes produce spermatozoa of simplified struc-
ture. The spermatozoa of these fish do not posses an acrosome (Stanley 1969), although
in some species there are structures which suggest that in the past their spermatozoa
possessed such an organelle (Mattei and Thiaw 1993). Moreover, activity of acrosin in
most of fish species has not been detected (Ciereszko et al. 1994). Acipenseriformes are
an exception among teleosteans in that their spermatozoa have acrosomes and active
acrosomal enzymes (AS, acrosin, β-NAGase). Changes in the structure of a male gamete
are accompanied by a specific structure of an egg cell in fish. A fish oocyte possess micro-
pyle, special channel through which spermatozoa can fuse with oocytes without having
to penetrate their outer membranes. On the one hand, as claims Jamieson (1991), this
model of gametes in fish shaped during their evolution has led to the presence of a single
micropyle, which prevents polyspermy. On the other hand, it caused loss of acrosomes.
Gametes of Acipenseriformes may represent an intermediary stage in evolution, as the
spermatozoa of these fish posses an acrosome (Jamieson 1991), but the surface of oocytes
typically has from 4 to 12 micropyles (Cherr and Clark 1984). Because of some differences
in the structure of gametes, the biochemistry of teleost fish semen has been an object
of the research carried out at the Department of Gamete and Embryo Biology, Institute
of Animal Reproduction and Food Research of the Polish Academy of Sciences. Recently,
very interesting findings have been made, namely the spermatozoa of trout have been
demonstrated to contain an acrosomal enzyme such as β-NAGase, although no activity of
another acrosomal enzyme. i.e. arylsulfatase has been observed (Sarosiek et al. 2008).
Biochemical research is essential for cryopreservation of semen, either to create
a gene bank for rainbow trout or to preserve endangered fish species, such as Acipenser-
iformes. Semen cryopreservation methods have become quite widespread, especially in
reproduction of farm animals. However, in respect of freezing fish semen, the methodol-
ogy is extremely difficult because the optimal freezing parameters must be selected indi-
vidually for each fish species (Medeiros et al. 2002). The biology of fish gametes remains
poorly recognized, therefore obtaining a satisfactory result of cryopreservation needs
multiple preliminary tests involving different variants of cryoprotectants. Cryoprotect-
ants are added in order to protect spermatozoa from the adverse effect of a fast freez-
ing rate. Cryoprotectants protect cells from dehydration, which appear during freezing.
They also stabilize structures of enzymatically active proteins. At the same time, cryo-
protectants should be characterized by low toxicity towards cells (Muchlisin 2005). In
228
Effect of cryoprotectants on activity of some enzymes and motility parameters of spermatozoa…
this paper, we have determined the effect of different cryoprotectants on the activity of
acid phosphatase, β-N-acetylglucosaminidase and arylsulfatase in seminal plasma and in
spermatozoa of rainbow trout and Siberian sturgeon. These two fish species have been
selected for the study because they represent two different types of gametes: trout has
spermatozoa without acrosomes while sturgeon has ones with this organelle.
Apart from the influence of cryoprotectants on the activity of the above enzymes,
their effect on the motility parameters of spermatozoa was assessed. Using the compu-
ter-assisted sperm analysis (CASA) software, researchers can determine objectively the
impact of many factors on the motility parameters of spermatozoa (Rurangwa et al.
2004). This is an extremely important aspect in our cognition of animal reproduction
process. The available literature provides very little information on such problems as the
effect of heavy metals on the motility of spermatozoa (Kime et al. 1996, Rurangawa et
al. 1998, Dietrich et al. 2010); likewise, we lack experimental results establishing the ef-
fect of cryoprotectants of the motility of spermatozoa of trout and sturgeon. This paper
provides us with the data regarding the effect of several cryoprotectants, such as DMA,
DMSO, methanol and glycerol, on the sperm motility of these fish species.

Semen of rainbow trout was obtained in April 2009 at the Salmonid Fish Breeding Sta-
tion in Rutki-Żukowo. Semen from each individual was sampled into separate tubes. Sam-
ples contaminated with blood, urine or faeces were discarded. Once transported to the
Department of Gamete and Embryo Biology, Institute of Animal Reproduction and Food
Research of the Polish Academy of Sciences in Olsztyn, at the temperature of +4oC, it was
centrifuged (10 min. 8000xg) in order to separate spermatozoa from semen plasma. The
sperm pellets and the seminal plasma obtained after the centrifugation of 8 ml of semen
from each male separately (n=5) were stored at -79oC for biochemical assays. Seminal
plasma and sperm pellets were thawed at room temperature, after which the spermato-
zoa were extracted for an hour in 5 ml 0.02 M Tris HCl, pH 7.6. The extract of spermatozoa
thus prepared was centrifuged twice (10 min. 8000xg) and used for biochemical assays.
Semen of Siberian sturgeon was obtained in Gryfino (Poland) using a catheter, having
previously stimulated the fish with Ovopel. The semen for analyses was transported to our
laboratory on ice, at the temperature of +4oC. Next, the milt for biochemical tests was
subjected to centrifugation (10 min. 8000xg) in order to separate plasma from spermato-
zoa. Afterwards, the biological material was frozen at -79oC for storage until proper analy-
ses. Afterwards, the spermatozoa thawed at room temperature were mixed with 0.02 M
Tris HCl, pH 7.6 and extracted for one hour. In the seminal plasma thus obtained and in the
supernatans of extracts from rainbow trout’s and Siberian sturgeon’s spermatozoa, the
enzymatic activities were determined with and without appropriate cryoprotectants.
Apart from the above, for each enzyme separately its activity was determined with an
addition of an appropriate inhibitor (negative control). The activity of acid phosphatase
(AcP) was measured using as a substrate 5 mM p-nitrophenyl posphate (disodium salt)
in 20 mM citrate buffer, pH 5.0 (Glogowski et al. 1996). In order to measure the activity
of β-N-acetylglucosaminidase, 0.5 mM p-nitrophenyl β-N-glucosamine in 0.1 M citrate
buffer, pH 5.0, was used (Jauhiainen and Vanha-Pertulla 1986). The activity of arylsulfa-
229
tase (AS) was measured using 0.02 M p-nitrocatechol sulphate in 0.5 M of acetic buffer,
pH 4.9 (Yang and Srivastava 1974). Enzymatic activities were determined in three repli-
cates. Solutions of 10% DMA (dimethyl acetamide), 10% DMSO (dimethyl sulphoxide),
10% methanol and 30% glycerol, 0.9 M sucrose, 0.001 M ammonium molybdate, 0.001
M AgNO3 and 1 M acetamide were prepared by dissolving the reagents in deionized
water. The above solutions were mixed with seminal plasma and spermatozoal extracts
of the two analyzed fish species in a 1 : 1 ratio. The enzymatic activity in seminal plasma
and extracts of spermatozoa was determined after 0, 3 and 14 hours of incubation with
the above solutions at +4oC.
Analysis of sperm motility (CASA) was carried out with use the equipment manufac-
tured by IH Medical. Fresh semen collected from 3 individuals of trout and 3 individuals
of sturgeon were mixed at a 1 : 1 ratio with immobilizing buffer (Morisawa for trout and
IMB for sturgeon), containing 10% DMA, 10% DMSO, 10% methanol and 30% glycerol
respectively (the final concentrations of all the analyzed reagents were half the above
values). The final concentrations of the above reagents correspond to the concentra-
tions used for cryopreservation. In order to activate motion, Biliard’s solution for trout
and Jahnichen’s solution for sturgeon were used, both supplemented with 0.5% of al-
bumin. The following parameters of the motility of spermatozoa were analyzed; VCL
(curvilinear velocity; μm/s), VSL (straight-line velocity; μm/s), VAP (average path velocity;
μm/s), LIN (linearity; %), STR (straightness; %), WOB (wobble; %) ALH – (lateral head am-
plitude; μm), BCF (beat cross frequency; Hz), MOT (percentage of motile spermatozoa;
%), Progressive (percentage of spermatozoa making progressive motion; %). The control
sample consisted of semen mixed with IMB without any additives.
The statistical analysis was performed using GraphPad Prism and applying one-factor
analysis of variance ANOVA for paired samples. Differences between particular experi-
mental variants were determined with post-hoc Tukey’s test.
Results
Rainbow trout
The activity of acid phosphatase in trout’s seminal plasma was too low for analyzing
the influence of cryoprotectants. In respect of acid phosphatase extracted from sperma-
tozoa, among the four analyzed substances, DMA and glycerol caused a 22 and 17% in-
crease in the activity of this
enzyme, respectively (Fig.
1). Addition of the other
cryoprotectants did not pro-
duce statistically significant
differences. No significant
differences were observed
in the activity determined
after 0, 3 and 24 h.
Analysis of the effect of
cryoprotectants on the ac- Figure 1. Effect of cryoprotectants on acid phosphatase activity present in
tivity of β-NAGase present extracts of rainbow trout’s spermatozoa.
230
Figure 2. Effect of cryoprotectants on β-NAGase activity present in rainbow

trout’s seminal plasma (A) and sperm extracts (B).
in trout’s seminal plasma

showed that DMA and
DMSO caused a significant
decrease in the activity of
this enzyme, by 18 and
11%, respectively. Addition
of 1 M acetamide, β-NA-
Gase inhibitor, led to a 35%
decrease the activity of the
enzyme (statistically sig-
nificant, Fig. 2). Although,
unlike acid phosphatase,
certain differences in the
activity of β-NAGase were
noticed depending on the
time of incubation with
a cryoprotectant. Moreo-
ver, the influence of a dose
of DMA and DMSO was
determined; consequently,
it was demonstrated that
5% concentration of either
of the cryoprotectants did
not lower the activity of β-
NAGase, whereas the con-
centration of 20% of DMA
decreased the activity of
β-NAGase by a significant
Figure 3. Effect of cryoprotectants on rainbow
value of 20%. The same en-
trout’s sperm motility parameters.
zyme present in spermato-
231
zoa of rainbow trout was inhibited by DMA and DMSO, which was manifested as its low-
er activity, whereas methanol caused increase in the activity of the analyzed enzyme.
Furthermore, the effect of the concentration of DMA and DMSO on the activity of
β-NAGase was assessed, and the results showed dose-dependent relation. In respect of
DMSO, similar values of the inhibition of β-NAGase were observed. 2 M concentration
of acetamide depressed the activity of β-NAGase by 74%, whereas at 1 M concentration
of acetamide, the inhibition reached 59% and 0.5M concentration inhibited the activity
of this enzyme by 42%.
Effect of cryoprotectants on the motility of rainbow trout’s spermatozoa – CASA

When analyzing the motility parameters of spermatozoa in the presence of different
cryoprotectants, it was demonstrated that addition of methanol and glycerol caused sta-
tistically significant decreasing in most of these parameters (Fig. 3). Additionally, it was
revealed that DMSO delayed the onset of spermatozoal motion by 5-6 seconds, whereas
DMA showed no influence of sperm motility. Glycerol caused a nearly complete inhibi-
tion of the motility of spermatozoa, which was most probably due to the high osmotic
pressure of the solution. Nonetheless, diluting the samples incubated with glycerol did
not result in any improvement of the sperm motility parameters.
Sturgeon
Addition of cryoprotectants to sturgeon’s sperm extracts did not lower the activity
of acid phosphatase (the activity of this enzyme in seminal plasma was too low to carry
out the tests, analogously to that of trout). Addition of an inhibitor of acid phosphatase,
such as ammonium molyb-
date, depressed the activity
of the enzyme by ca 75%
compared to the control
(Fig. 4).
β-NAGase from stur-
geon’s seminal plasma
proved to be very sensitive
to addition of cryoprotect-
ants, particularly that of
Figure 4 Effect of cryoprotectants on acid phosphatase activity
DMSO and DMA, which present in Siberian sturgeon’s sperm extracts.
were found to depress its
activity by around 50% (Fig. 5A). The influence of DMA depended on a dose: 20%
concentration of DMA caused 49% inhibition of the activity of β-NAGase, whereas 5%
DMA inhibited the activity of this enzyme by 28%, Similar relationships were demon-
strated in the case of DMSO: 20% concentration of this cryoprotectant caused 60%
inhibition of the activity of this enzyme whereas 5% DMSO inhibited its activity by
29%. The other cryoprotectants, except methanol, likewise caused decreasing β-NA-
Gase activity. Furthermore, it was demonstrated that this enzyme was the most active
after 3 hours of incubation with any of the tested cryoprotectants. At the ‘0’ time, the
232
activity was the lowest. The negative control, composed of a sample with addition of
β-NAGase inhibitor (acetamide) showed that the activity of this enzyme significantly
decreased in the presence of the inhibitor. 2 M concentration of acetamide caused
a loss of 83% of the enzymatic activity, while at 0.25 M concentration the loss was
51%. The activity of β-NAGase present in sturgeon’s spermatozoa was also inhibited
in the presence of DMA and DMSO. 1 M acetamide caused an up to 88% inhibition of
the activity of this enzyme (Fig. 5B).
Addition of any of the tested cryoprotectants to sturgeon’s seminal plasma caused
small (10-16%) but statistically significant decrease in the activity of arylsulfatase. No
differences were observed between the effect of particular cryoprotectants (Fig. 6), in
Figure 5. Effect of cryoprotectants on β-NAGase activity present in Siberian sturgeon’s

seminal plasma (A) and sperm extract (B).
Figure 6. Effect of cryoprotectants on arylsulfatase activity present in Siberian sturgeon’s

seminal plasma (A) and sperm extracts (B).
contrast to the results of our analysis of the enzyme derived from sperm extracts, where
addition of the cryoprotectants did not cause any significant lowering of the activity of
arylsulfatase. Regarding seminal plasma and sperm extracts, no differences were found
in the activity of AS depending on the incubation time.
233
Effect of cryoprotectants on the motility of spermatozoa of Siberian sturgeon – CASA

When analyzing the effect of cryoprotectants on the motility of spermatozoa of stur-
geon, it was only glycerol that was demonstrated to have caused significant decrease of
the sperm motility param-
eters (Fig. 7). Moreover,
after addition of DMA and
methanol the value of Pro-
gressive decreased.
Discussion
The biological value
of frozen semen is lower
than that of fresh one,
which has been evidenced
for both mammals (Holt
2000, Watson 2001) and
fish (Rana 1995, Babiak
et al. 1998, Glogowski et
al 2001). In this paper we
show that the activity of
enzymes present in seminal
plasma and spermatozoa
of teleost fish did not un-
dergo significant changes
in the presence of cryopro-
tectants, except the activity
of β-NAGas. This enzyme,
whether in seminal plasma
or in sperm extracts, proved
to be sensitive to addition of
two cryoprotectants, such
as DMA and DMSO. This is
a very interesting finding if
Figure 7. Effect of cryoprotectants on Siberian
we compare it to the data sturgeon’s sperm motility parameters.
obtained by Sarosiek et al.
(unpublished data), where the activity of β-NAGase in spermatozoa and oocytes of trout
and sturgeon was inhibited. It was demonstrated that the percentage of fertilized eggs
was drastically lowered in both species. Thus, the role of this enzyme during fish ferti-
lization is extremely important. Determination of the effect of cryoprotectants on the
activity of enzymes, and particularly β-NAGase, implies that similar research can be use-
ful for making a choice of a suitable cryoprotectant while preparing a cryopreservation
protocol for the species for which such procedures have not been described.
Motility of spermatozoa is a measure of their quality and ability to fertilize egg cells
(Lahnsteiner et al. 1998). The experiment run with the CASA system, which determines
234
parameters of sperm motility, enabled us to prove that addition of glycerol and methanol
caused decreasing in sperm motility parameters for trout. Therefore, it is most probable
that glycerol as well as methanol have a nearly toxic effect to the motion apparatus of
trout’s spermatozoa. Our analysis of the sperm motility parameters of sturgeon’s sperma-
tozoa with and without cryoprotectants showed that these parameters tend to decrease
in the presence of cryoprotectants but the differences were not significant statistically
when DMSO, DMA and methanol were applied (Fig. 7). However, it was found out that
in the presence of DMA and methanol, the percentage of motile spermatozoa (MOT) in-
creased. It was also discovered that all the applied cryoprotectants caused decreasing in
the values of progressive motion (Progressive). Among all the tested cryoprotectants, it
was only glycerol that caused an evident, statistically significant, decreasing in the motil-
ity parameters, which was most probably due to considerable viscosity of the solution.
However in the tests aiming at viscosity reduction, some improvement in the sperm mo-
tion parameters was observed, although the results were not satisfactory (unpublished
data). Glogowski et al. (2002) obtained good results when freezing sturgeon’s semen with
methanol as a cryoprotectant. The results obtained in our experiment confirm that metha-
nol does not affect both the activity of the tested enzymes nor the motion apparatus of
spermatozoa. Considering the above results, it can be concluded that DMA and methanol
show the weakest influence on the motility of sturgeon’s spermatozoa. Nonetheless, the
negative effect that DMA has on the activity of β-NAGase eliminates this compound from
being a potential cryoprotectant used for freezing sturgeon’s semen.
Acknowledgements
The experiment completed a part of the research project of the Ministry for Science
and Higher Education no N308 018 31.
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Effectiveness of Ovaprim in
controlled reproduction of some
cyprinid fish species. Part I:
latency time and synchronization
of ovulation
Daniel Żarski1, Katarzyna Targońska1,
Sławomir Krejszeff1, Katarzyna Palińska2,
Sebastian Piszczała3, Dariusz Kucharczyk1
1
Department of Lake and River Fisheries, University of Warmia
and Mazury in Olsztyn, Oczapowskiego 5, 10-957 Olsztyn, Poland,
tel. +48 895234436, fax + 48 895233969,
e-mail: danielzarski@interia.pl
2
Department of Zoology, University of Warmia and Mazury in
Olsztyn
3
Knieja Fish Farm
Abstract
The study concerned the effectiveness of two commercial spawning agents (Ovopel
and Ovaprim) in reproduction of ide, dace and common nase. The tests involving ide and
dace were carried out in a strictly controlled environment, whereas common nase was
studied under hatchery conditions, with no control over environmental factors. The re-
sults suggest that both of the applied preparations were highly useful for reproduction
of these species. When fish were administered Ovaprim, a higher percentage of ovulat-
ing females was observed among all the three species. The latency time in ide and dace,
which received this preparation was longer and spawning much more prolonged com-
pared to Ovopel. With respect to common nase, where daily fluctuations in temperature
appeared, the latency time was the same after administration of either of the prepara-
tions (51h) but oocytes were obtained much later when Ovopel was administered. The
results clearly indicate that Ovaprim was more effective than Ovopel.
Introduction
Over the past decades, due to climatic changes, pollution and transformation of the
natural environment, the ecological condition of inland waters has greatly deteriorated.
237
As a result, habitats of many endemic fish populations have undergone degradation.

This strongly affected rheophilic cyprinids, whose life cycle is strictly dependent on dif-
ferent biotopes (Mann 1996, Penczak 1998). Lack of suitable spawning grounds or mi-
crohabitats for offspring can lead to a serious limitation of the abundance of these fish
or even extinction of some populations. Consequently, it is necessary to work out resti-
tution programmes, in which reproduction under controlled conditions is an essential
component (Philippart 1995, Kucharczyk 2002).
Completely controlled reproduction of wild fish populations requires that previous
hormonal stimulation be performed (Kucharczyk 2002, Szabo et al. 2002, Heyrati et al.
2007, Krejszeff et al. 2008), therefore, in the recent years, many spawning agents have
appeared on the market (Yaron 1995, Brzuska 2000, Szabo 2003, Brzuska 2005). The
applied hormonal preparations may affect in a variety of ways the synchronization of re-
production, including species related to one another. Thus, it is important to determine
experimentally the type and rates of spawning agents which ensure the highest possible
effectiveness (Kucharczyk 2002, Yaron et al. 2009).
In induced reproduction of cyprinid fish, one of the most commonly used prepara-
tions is Ovopel (Unic-Trade). It is a Hungarian preparation, which contains mammalian
gonadotropin-releasing hormone (GnRH) analogue and a dopamine antagonist – meto-
clopramide (Horvath et al. 1997). It is commercially available in the form of pellets,
which should be homogenized in a sterile solution of saline (0.9% NaCl). The suspension
thus obtained is administered as injections. Another preparation, which has recently
come into use in aquaculture of cyprinids is Canadian Ovaprim (Syndel) – an agent
which is a complex of salmon GnRH analogue and a dopamine antagonist domperidone
(Brzuska and Adamek 1999). This preparation is sold in the form of oily liquid, ready for
administration.
The objective of this paper has been to compare the effectiveness of the two com-
mercial preparations, Ovopel and Ovaprim, in stimulation of ovulation in some rheophil-
ous cyprinid species.

Spawners of ide (mean weight 167±9g) were captured in the Marózka River (north-
eastern Poland), while those of dace (mean weight 411±21g) were supplied by the Fish
Farm in Janowo near Szczytno. The experiments on ide and dace were completed in
a hatchery of the Department of Lake and River Fisheries of the University of Warmia and
Mazury in Olsztyn, whereas common nase was studied at the Station of the Department
of Applied Ecology of the University of Łódź in Tresta. During the experiments, females and
males were kept separately in 1,000 dm3 tanks. Reproduction of ide and dace was carried
out using thermal stimulation according to the protocol described by Kucharczyk et al.
(2008a), whereas spawners of common nase were kept in water of natural thermal con-
ditions. The range of temperatures during the reproduction of common nase varied be-
tween 7.4 and 13.4oC. The photoperiod for ide and dace was 12 h (12L : 12D). During the
reproduction of common nase, the daylight was not controlled (natural photoperiod).
Reproduction of all the three species was conducted using two hormonal prepara-
tions: Ovopel and Ovaprim. Ovopel contains 18-20 μg GnRH analogue [(D-Ala6, Pro9-
238
Effectiveness of Ovaprim in controlled reproduction of some cyprinid fish species.…
Net)-mGnRH] and 8-0 mg metoclopramide (Horvath et al. 1997). Ovaprim contains 20

μg GnRH analogue [(D-Arg 6 , Pro9-Net)- sGnRH] and 10 mg domperidone in 1 ml solu-
tion (Peter et al. 1993). In respect of ide and dace females, two injections of Ovopel (in
24 h intervals) and one injection of Ovaprim were administered. Common nase females
received one injection (Tab. 1). The onset of ovulation in ide was checked for the first
time 36 hours after the last injection and in dace and nase after 48 hours. Afterwards, it
was checked every 3-4 hours.
During the manipulations, the spawners were anesthetized in a solution of 2-phe-
noxyethanol (Sigma-Aldrich, Germany) of the concentration equal 0.5 cm3 dm-3. All the
manipulations, including the eggs stripping, were performed according to the method-
ology described by Targońska et al. (2008) for dace. During the study, the number of
spawned females and the time between the resolving injection and ovulation (latency
time) were recorded.
Table. 1. Types and doses of hormonal agents administered in controlled reproduction
of ide, dace and common nase
Ide and dace

Group Common nase
1st injection 2nd injection
1 cm3 kg-1 1 cm3 kg-1 1 cm3 kg-1

Control
(0.9 % NaCl) (0.9 % NaCl) (0.9 % NaCl)
0.2 granules kg-1 1 granule kg-1 1 granule kg-1

Treatment 1
(Ovopel) (Ovopel) (Ovopel)
0.5 cm3 kg-1 0.5 cm3 kg-1

Treatment 2 -
(Ovaprim) (Ovaprim)
Results
In all the three species, a positive effect of both spawning agents on ovulation was
observed. Eggs were not obtained from any female from the control groups. The results
recorded during the experiment are presented in Table 2.
Ovulation in dace females was achieved between 48 and 72 hours after the resolving
injection. When Ovopel was administered, this time period was shorter and spawning
was finished 50 hours after the last injection. Ovaprim caused desynchronization of
females becoming ready to spawn – eggs from the last female were obtained 72 hours
after the last injection. In the group receiving Ovoopel, eggs were obtained from 90% of
the females. Administration of Ovaprim caused ovulation in all the fish.
Regarding dace, the latency time was within the ranges of 36-38 and 38-48 hours
after the administration of Ovopel and Ovaprim, respectively. In the group treated with
Ovopel, ovulation was certified in 90% of the females. In the group stimulated with
Ovaprim, eggs were obtained from all the females. No synchronization in ovulation was
observed in the latter group.
239
Table 2. Results obtained during controlled reproduction of ide,

dace and common nase using Ovopel and Ovaprim
Control group Ovopel Ovaprim

Species Dace
Number of females 9 10 10
Percentage of ovulating females
0 90 100
(%)
Latency time (hours) Not applicable 48 - 50 48 - 72
Synchronization of ovulation Not applicable Yes No
Species Dace
0 90 100
(%)
Latency time (hours) Not applicable 36-38 38-48
Synchronization of ovulation Not applicable Yes No
Species Common nase
0 89 100
(%)
Latency time (hours) Not applicable 51-75 51
Synchronization of ovulation Not applicable No Yes
In the particular treatment groups comprising common nase, the latency time was
varied. The administration of Ovaprim caused ovulation in all females 51 hours after the
injection. In respect of the fish which received Ovopel, eggs were obtained between 51
and 75 hours. The last female which ovulated in response to Ovopel spawned spontane-
ously in the tank over 75 hours after the injection. Ovopel affected successful spawning
in 89% of the females.
Discussion
The latency time is an important parameter in controlled fish reproduction. What is
of special importance is the time period during which ovulation of high quality oocytes
is expected. In practice, this parameter manifests itself in the effectiveness of fish stock
material production. Obtaining small amounts of eggs from individual females is la-
bour-intensive and stressful for fish. Besides, it increases the costs of production and
poses a risk of losing some of the spawners. Another problem appears during incuba-
tion of eegs when a larger number of incubators needs to be used. It also results in pro-
longed hatching of larvae, which may affect size heterogeneity of reared fish and lead,
for example, to cannibalism (Kwiatkowski et al. 2008, Żarski et al. 2008a). The latency
time depends on several factors, such as species, maturity of females, photoperiod and
temperature, including daily fluctuations in temperature and thermal acclimation rate
(Żarski et al. 2010, Targońska et al. – unpublished data). The temperature of water from
which spawners are caught also matters.
240
Synchronization of ovulation using hormonal stimulation has been practiced for many
years. Initially, it relied on introduction to a fish body exogenous gonadotropins with the
homogenate of pituary glands of fish (Drori et al. 1994, Yaron 1995). However, due to vari-
able quality of such hypophysate, research has been undertaken on other agents. GnRH
analogues have been used successfully in controlled fish reproduction, but their effective-
ness in many species of cyprinid fish depends on simultaneous administration of com-
pounds which are dopamine antagonists (Peter and Yu 1997, Zohar and Mylonas 2001).
For this purpose, complex preparations have been made, which - apart from being highly
successful - are available in easily administered form of granules or liquid. The two stimu-
lating agents tested in this study belong to such hormonal preparations.
Different latency periods following administration of hormonal agents have been
reported elsewhere (e.g. Yaron 1995, Brzuska 2000, 2005). However, the differences
most often concerned a different response of fish organisms to hypophysate and to go-
nadotropin-releasing hormone analogues, which act on different levels (gonads and the
pituary gland, respectively). In the present study, two agents affecting fish at the same
level, although different in composition, have been tested. Different response expressed
in the latency time and caused by administration of different GnRH analogues has been
recorded for carp Cyprinis carpio (L.) (Brzuska 2006). A similar dependence has also been
observed by Jamróz et al. (2008), who found the best synchronization of ovulation in
female ide having administered Ovopel (36h), in contrast to the group stimulated with
Ovaprim, in which ovulation was observed between 36 and 44 hours after the resolv-
ing injection. A longer latency time after administration of Ovaprim in comparison to
Ovopel has also been found in a study on asp Aspius aspius (L.) (Żarski et al. 2008a). An
similar dependence, as in the above reports, has been determined for dace, which was
for example confirmed by Kucharczyk et al. (2008a).
With respect to common nase, the time elapsing from the injection to production
of gametes was 11 hours longer than cited by Szabo et al. (2002), who likewise used
a GnRH analogue with dompamine antagonist but maintained constant temperature
of water during their experiment (12oC). Moreover, for the group of fish which were
administered Ovopel, the spawning period was prolonged by another 24 hours, which
was most probably related to the unstable thermal conditions, because the tests were
conducted in water of natural temperatures. It is a known fact that variations in tem-
perature can either delay or accelerate (depending on a fish species) the moment when
females were ready to spawn (e.g. Bromage et al. 2001, Davies and Bromage 2002, An-
gius and Cańavate 2005). Therefore, synchronization of the moment of ovulation after
administration of Ovaprim seems to suggest that this agent was much more effective in
the controlled reproduction of common nase. In that case, Ovopel resulted in the prolon-
gation of latency time. The results achieved in this study indicate that the administered
hormonal preparations are highly useful for the controlled reproduction of rheophilic
cyprinid fish. Although Ovaprim administered to ide and dace caused desynchronization
of ovulation in individual females, it should be mentioned that it had the most successful
effect (100% ovulating females). High effectiveness of this agent has been additionally
verified by the tests on reproduction of common nase under unfavourable thermal con-
ditions. It should be added here that the differences in the action of both preparations
241
may be attributable to the different types of analogues they contain. Podhorec and
Kouril (2009) claim that the salmon form of GnHR appears naturally in cyprinids. There-
fore, the salmon GnRH analogue contained in Ovaprim (more closely similar in structure
to the natural hormone than the mammalian analogue) most probably affects the or-
ganisms of cyprinid fish more effectively than the mammalian analogue. Thus, a much
smaller dose (nearly four-fold less) of the analogue causes similar and sometimes even
superior effects on reproduction in this group of fish (Targońska et al. 2010). However, it
is impossible to exclude with absolute certainty a significant effect of different types of
dopamine antagonists. Consequently, more detailed studies on induced reproduction of
cyprinid fish involving this group of hormonal agents are needed.
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243
Effectiveness of Ovaprim in
controlled reproduction of some
cyprinid fish species.
Part II: gametes quality
Katarzyna Palińska1, Daniel Żarski2,
Sławomir Krejszeff2, Katarzyna Targońska2,
Sebastian Piszczała3, Dariusz Kucharczyk2
1
Department of Zoology, University of Warmia and Mazury in
Olsztyn, e-mail: katarzyna.palinska@uwm.edu.pl
2
Department of Lake and River Fisheries, University of Warmia and
Mazury in Olsztyn
3
Knieja Fish Farm
Abstract
The reported study deals with the effect of application of two hormonal preparations
(Ovopel and Ovaprim) on the biological quality of gametes obtained during induced
reproduction of ide and dace. During the experiments, controlled reproduction of these
two fish species was carried out. The biological quality of gametes was determined
based on the survivability of embryos in the eye-egg stage. The results indicate that
Ovaprim is much more useful than Ovopel. This conclusion holds true for the spawning
out-of (72.7% live embryos for ide and 61.2% for dace) and during the natural spawning
season (77.5 and 60.5% for ide and dace, respectively).
Introduction
A decline in the abundance of fish, observed for many years now, either caused by
angling or fishing exploitation (e.g. Penczak 1998, Skrzypaczak 2000), necessitates ac-
tive protection. This problem largely concerns rheophilic cyprinid fish (Augustyn 2004).
Among the most important tools serving active protection of endangered fish species
is aquaculture (Philippart 1995, Poncin and Philippart 2002). Studies carried out for many
years now have contributed to the elaboration of complex procedures applied to rearing
fish stock material, which include such aspects as the welfare of the natural environment
and the economic profitability of fish restitution programmes (Kucharczyk et al. 1008b,
245
Kupren et al. 2008, Targońska et al. 2008). Controlled reproduction has become the base
for restitution programmes and directly affects the quality of produced fry. Moreover, it
enables us to gain full control over the moment when offspring is produced because re-
production can be carried out out-of the spawning season, which may shape the effective-
ness of fish restocking (Cowx 1994, Kucharczyk et al. 1999, Kucharczyk 2002).
The volume of production of rheophilic cyprinid fish stocking material has increased
over ten-fold between 1995 and 2005, reaching nearly 83 tons (Wojda 2004, Mickiewicz
et al. 2007). The highest percentage was made up by ide, whose fish stock material pro-
duction first reached 99% of the total share of cyprinid rheophilic fish in the total fish re-
leases to open waters. The data of 2005 indicate that the percentage of ide fell to 70%,
owing to a much larger production of the other species of rheophilic cyprinid fish.
The effectiveness of the fish reproduction procedure under controlled conditions can
be estimated according to several parameters, like the ovulation rate, latency time and
survivability of spawners, which is a result of the applied handling procedures. How-
ever, the most important parameter is the survivability of embryos, which also indirectly
characterizes the quality of gametes, and especially oocytes (Bobe and Labbe 2010). In
respect of cyprinid fish, the most frequently applied parameter is the determination of
survivability of embryos at the moment when pigment appears in the eyes, i.e. in the
eye-egg stage (Kucharczyk 2002). Regarding the eggs of rheophilic cyprinid fish, this is
the earliest moment when dead embryos can be objectively distinguished from live ones.
Besides, this moment is easy to capture and it is possible to calculate the percentage of
live embryos macroscopically.
The objective of the paper has been to determine the effect of the applied spawning
agent on the biological quality of gametes of two rheophilic cyprinid fish species, under
controlled reproduction conditions, during and out-of the spawning season.

The material consisted of gametes of ide and dace obtained via controlled reproduction
during and out-of the spawning season. Ovulation was induced through administration
of two commercial spawning agents: Ovopel and Ovaprim. Ide spawners (average weight
170 g) were caught in the Marózka River, whereas dace spawners (average weight 450g)
originated from the Fish Farm in Janowo near Szczytno. The reproduction of both species,
during and out-of the spawning season, was carried out according to the methodology
described by Kucharczyk et al. (1999) for ide. The hormonal stimulation of ide and dace
was performed with the doses given by Żarski et al. (2009). Thus, Ovopel was administered
in two doses (0.1 and 1 granule kg-1) whereas Ovaprim was given as a single injection (0.5
cm3 kg-1). These doses have been, until present, the smallest effective doses established ex-
perimentally and commonly applied in cyprinid fish reproduction (Kucharczyk et al. 2008b,
Targońska et al. 2010). One pellet of Ovopel contains 18-20 μg of gonadoliberine (GnRH)
[(D-Ala6, Pro9-Net)-mGnRH] analogue and 8-10 mg metoclopramide (Horvath et al. 1997).
Ovaprim, sold as a liquid, contains 20 µg GnRH [(D-Arg 6 , Pro9-Net)- sGnRH] analogue and
10 mg domperidone in 1 ml solution (Peter et al. 1993). Prior to the hormonal stimula-
tion, fish were randomly divided according to the gender and hormonal treatment. Once
they were found to ovulate, eggs were sampled into dry plastic containers. Next, a sample
246
Effectiveness of Ovaprim in controlled reproduction of some cyprinid fish species…
of eggs was taken from each female (about 150-200 grains) and placed on Petri’s dishes
in three replicates. The eggs samples were fertilized with a pooled semen (0.05 cm3) ob-
tained from the males which had received an injection of the same hormonal agent. After-
wards, the gametes were activated with water. Incubation was carried out in a close water
circulation system equipped with an aerator and the equipment which enabled to control
the temperature at 0.1oC accuracy. The spawn of dace was incubated at 12oC and that of
ide at 14oC. The moment pigment appeared in the embryos’ eyes (in the eye-egg stage),
the survivability of embryos was determined.
Statistical differences between the groups were analyzed with analysis of variance
(ANOVA), and next using post-hoc Tukey’s test (P<0.05). Before the analysis, all the data
expressed in per cent were arc-sin transformed.

In the hatchery practice, during the whole fish culture cycle, many factors affect
the effectiveness and consequently the economic profitability of fish stocking mate-
rial production (under controlled conditions). These factors pertain to the biotechnical
procedures of reproduction and rearing of fish fry. As for reproduction, the origin of
spawners is essential (Krejszeff et al. 2009). If obtained from the natural environment,
it is important how spawners were captured (Kujawa et al. 2006), because this may
have some influence on their fitness in captivity. Other factors which determine the
production success are: the method and conditions under which spawners were kept
in a hatchery (Kujawa et al. 1999), the rate of thermal acclimation (Żarski et al. 2010,
Cejko et al. – unpublished data, Targońska et al. – unpublished data), the moment when
reproduction is carried out versus the natural spawning season (Kucharczyk et al. 1999,
Targońska-Dietrich et al. 2004, Kucharczyk et al. 2008a) as well as the type and dose of
applied spawning agent (Brzuska 2006, Kucharczyk 2002, Jamróz et al. 2008, Krejszeff
et al. 2008, 2009, Kucharczyk et al. 2008b, Targońska et al. 2008).
The effects of hormonal stimulation of ide, as evidenced in the present experiment
(Tab. 1), confirm the usability of Ovopel in controlled reproduction of this species.
Moreover, the results were close to the ones reported by Kucharczyk et al. (1999), who
obtained 79.3 and 63.6% survivability of embryos respectively during and out-of the
spawning season. These authors achieved the best reproduction results after stimula-
tion with Ovopel, compared to the treatments with carp pituary homogenate (CPH)
and human chorionic gonadotropin (hCG). These results were also very close to the
ones achieved by Targońska-Dietrich et al. (2004) out-of the spawning season among
broodstock originated from fish ponds and cultured in a recirculation system (84.1 and
68.1% of embryo survivability, respectively). Noteworthy is the fact that after application
of Ovaprim, the survivability of embryos recorded in this study was higher than after the
application of Ovopel. When the fish reproduction was conducted out-of the spawning
season, lower survivability of embryos of both species was noted after the application of
Ovopel compared to the results obtained during the spawning season. When Ovaprim
was administered, the percentage of live embryos among female dace was slightly lower
during the breeding season (Tab. 1). However, it should be emphasized that reproduc-
tion run out-of the spawning season is associated with a considerable decline in the
247
quality of obtained gametes (Kucharczyk et al. 19999, Targońska-Dietrich et al. 2004,

Kucharczyk et al. 2008a). The relationships concerning the effectiveness of the tested
preparations were confirmed by the results reported by Kucharczyk et al. (2007) and
Jamróz et al. (2008). Higher survivability of embryos after Ovaprim stimulation has also
been found for asp Aspius aspius (L.) (Targońska et al. 2010).
Table 1. Survival of embryos of dace and ide at the eyed-egg stage obtained after controlled reproduction (performed
during and out-of the spawning season) with the application of two spawning agents: Ovopel and Ovaprim.
During spawning season Out-of the spawning season

Ovopel Ovaprim Ovopel Ovaprim
Species dace
Number of females 9 10 10 10
Survival of embryos at the eyed-
58,6 ± 5,6 b
77,5 ± 2,6 c
48,3 ± 4,5a
72,7 ± 2,7c
egg stage (%)
Species ide
Number of females 9 10 10 10
Survival of embryos at the eyed-
52,4 ± 3,1b 60,5 ± 2,3c 44,3 ± 3,8a 61,2 ± 3,1c
egg stage (%)
* Data in rows marked with different superscripts were statistically different (P<0.05).
The quality of gametes produced during controlled reproduction is affected by a

variety of factors. Research conducted for many years indicates that one such factor is
the hormonal stimulation, and the type or dose of a spawning agent influences con-
siderably the quality of oocytes and parameters of semen. Many scientific papers have
been devoted to the effect of administered hormonal preparations on the survivability
of embryos (e.g. Kucharczyk 2002, Brzuska 2006, Kucharczyk et al. 2008a, Targońska et
al. 2008). Most papers on cyprinid fish induced reproduction imply that preparations
containing active analogues of gonadotropin-releasing hormone (GnRHa) in combina-
tion with a dopamine antagonist (e.g. metoclopramide, domperidone) were more suc-
cessful. The effect of the type of a hormonal agent on the volume of obtained semen
and concentration of spermatozoa has been previously confirmed in different fish spe-
cies (Kucharczyk 2002, Targońska-Dietrich et al. 2004, Cejko et al. 2008). Administration
of Ovopel to cyprinid fish has produced much better results than stimulation of CPH or
hCG. In turn, the highest concentration of spermatozoa as well as their most superior
motility and biochemical parameters in asp have appeared as a result of administration
of Ovaprim rather than Ovopel (Cejko et al. 2008). A significantly higher percentage of
motile spermatozoa in semen from males injected with Ovaprim has also been deter-
mined in the case of ide (Jamróz et al. 2008). In this study, both males ad females (within
treatment groups) were stimulated with the same spawning agent. The effect such as
statistically better results may have been a synergistic effect of the positive influence
of Ovaprim on the quality of both oocytes and semen. Podhorec and Kouril (2009) re-
ported that the naturally occurring form of gonadoliberin in cyprinid fish is the salmon
form (pGlu1 – His2 – Trp3 – Ser4 – Tyr5– Gly6 – Trp7 – Leu8 – Pro9 – Gly-NH210). The analogue
found in Ovaprim is different from the natural form in the 6th and 10th position of the
248
Effectiveness of Ovaprim in controlled reproduction of some cyprinid fish species…
decapeptide chain, whereas mGnRHa (the mammalian analogue of gonadoliberin) is

different in positions 6, 7, 8 and 10 compared to sGnRHa (the salmon analogue of go-
nadoliberin). Therefore, it can be suspected that the sGnRHa form (the salmon analogue
of gonadoliberin) contained in Ovaprim, by being closer to the form naturally occurring
in cyprinid fish, has a positive effect on the course of final oocyte maturation and ovula-
tion. However, further studies are needed in order to achieve better understanding of
the mechanisms involved in the action of Ovaprim.
The results reported in this paper suggest that Ovaprim is more efficient than Ovopel.
The former agent has a positive influence on the ovulation of females (Jamróz et al.
2008, Żarskiet al. 2009) and on a statistically higher percentage of live embryos. Thus,
administration of Ovaprim may lead to production of much more fry and, in combina-
tion with an appropriately selected rearing method, it can considerably raise the profit-
ability of production of fish stocking material of rheophilic cyprinid fish.
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250
Differences in the efficiency of
controlled reproduction of carp
Cyprinus carpio L. of the German
line N, Hungarian line 7 and
Hungarian line 8
Elżbieta Brzuska
Polish Academy of Sciences, Institute of Ichthyobiology and Aqua-

culture,
Gołysz, 43-520 Chybie, Poland, tel. 0048 33 8533778 fax. 0048 33
8589292 e-mail: urszula.karosek@fish.edu.pl
Abstract
The paper contains the results of controlled reproduction of three breeding lines
of carp (German line N, Hungarian line 7 and Hungarian line 8) after stimulation with
Ovopel (1/5 pellet kg-1 body mass followed by 1 pellet kg-1 after 12 hours). Ovulation
occurred in 100% of females of line 8 and 80% of females of lines N and 7; it was only
among line 8 females that no cases of partial ovulation were noticed. The origin of the
females significantly (P≤ 0.05) conditioned the mass of the eggs obtained, both ex-
pressed in grams and as a percentage of the body mass of females. The highest means
of these traits characterizing the weight of spawn were determined for line 8 females
(1007.16g and 11.12%). They were significantly different (P≤0.05) from the means com-
puted for line N and line 7 females (720.59 and 9.72%, 709.16g and 7.15%, respec-
tively). The breeding line conditioned significantly (P≤0.05) two other features which
describe the quality of spawn, i.e. percentage of live embryos after 48 and 56 hours of
incubation. The lowest (75.82%) value of the mean of the least squares for the percent-
age of live embryos after 56 h incubation was recorded for line 7 females. The values
of the means of that feature for the two other lines oscillated around 82%, thus being
significantly different from that determined for line 7 females. Considering the values of
all the analyzed characteristics, calculated for the three lines of carp, it can be concluded
that the highest efficiency of reproduction was demonstrated by line 8 and the poorest
one – by line 7. Based on the results of the step regression calculations with the aim of
predicting the percentage of live embryos after 56 hours of incubation and according
251
to the distribution of the analyzed characteristics in a spatial arrangement, it has been

demonstrated that the most precise prediction for this feature was attainable for line N,
whereas the least precise one was achieved for line 8.
Introduction
The Institute of Ichthyobiology and Aquaculture of the Polish Academy of Sciences
in Gołysz possesses 19 breeding lines of carp (Białowąs 2003). These lines, different in
origin (Hungarian, Polish, Yugoslavian, French, German, Ukrainian, Lithuanian, and Is-
raeli), are valuable material for genetic (Irnazarow 1994, Iznazarow and Białowąs 1995,
Rakus et al. 2003) and immunological research (Kachamakova et al. 2006, Jurecka et al.
2009) of this commercially important fish species. This genetically variable herd of carp
fish, created over dozens of years of breeding and genetic studies carried out at the In-
stitute, is excellent material for Polish programmes designed to maintain genetic reserve
of breeding fish and to improve the culture of the Polish population of carp.
These lines have also been used as material for a long-term study carried out at the
Institute of Ichthyobiology and Aquaculture PAS in Gołysz on effects on reproduction of
carp originating from different areas. In these studies, results of controlled reproduction
of carp have been analyzed after stimulation with natural and synthetic preparations
(Brzuska 1991, 2004, 2006a,b, Brzuska and Ryszka 1990, Brzuska and Adamek 1997).
The Hungarian stimulant Ovopel, which is a complex of the mammalian GnHR ana-
logue (D-Ala6, ProNEt-mGnRH) and metoclopramide (Horvath et al. 1997), has been of-
ten used in experiments carried out in our institute, on the material composed of carps
from different breeding lines (Brzuska 2001, 2003, 2006a, 2010, Brzuska and Grzywac-
zewski 1999, Brzuska and Białowąs 2002, Cejko 2007). This article signals some progress
achieved during the next stage of our studies, carried out on genetically varied material
treated with Ovopel, the most commonly applied ovulation stimulant in fish breeding
practice of both cyprinid and other fish species.
The purpose of this study has been to demonstrate whether there are any differences
in the effects of controlled reproduction of three breeding lines of carp, of which two
had not been previously submitted to this type of study after Ovopel stimulation.

The experiment was run at the Institute of Ichthyobiology and Aquaculture of the
Polish Academy of Sciences in Gołysz. It comprised the data obtained during controlled
reproduction of carps originating from German line N (Bialowąs 2003), Hungarian line
7 (Białowąs 1999) and Hungarian line 8 (Białowąs 1998). Among these lines, neither
German line N nor Hungarian line 8 had been previously studied in terms of results of
reproduction obtained after stimulating females with Ovopel.
Thirty 8-year-old females, ten from each line, were taken for controlled reproduction.
The fish were selected from larger populations based on their external signs of maturity.
The body weight of the females ranged from 6.35 to 12.70 kg. The fish were transported
from the pond to a hatchery, where they were placed in three tanks, each containing 3
m3 of water of the temperature of 19oC (±1oC) After twelve hours of adaptation, stimu-
lated ovulation began. 1/5 of a pellet of Ovopel per 1 kg of weight was administered
252
Differences in the efficiency of controlled reproduction of carp Cyprinus carpio L. of the German…
to initiate ovulation, followed by 1 pellet per 1 kg b.w. to activate it. The time interval
between the intraperitoneal injections was 12 hours.
The monitoring of ovulation started 9 hours after giving the releasing injection
and continued every hour for the next 8 hours. Eggs obtained from each female were
weighed separately and then fertilized with pulled semen collected from 3-4 males,
which were stimulated to produce sperm with 1 pellet of Ovopel per 1 kg b.w. (intra-
peritoneal injection). Fertilized eggs weighing ~300 g from each female were incubated
separately in Weiss’s jars of the capacity of 3 l of water at the temp. 21oC (±1oC). After
12 hours of incubation, percentage of fertilization was calculated for each female, and
then after 24, 36, 48 and 56 hours of eggs incubation – percentage of live embryos was
assessed.
The data thus obtained underwent least squares analyses of variance (Harvey 1987),
completed according to the following formula:
Yij = α + ri + Wbcij + eij 200
g A
160
where: 120
80 D=812,31
40 line 8
α – theoretical general 0
-40
line 7
line N
mean -80
-120
-160
ri – effect of the ith ori- -200
gin (breeding line) % B % C

5 5
b – regression on body 4
3
D=9,33
4
3
D=94,90
2 2
weight of females 1
0
1
0
Wcij –body weight of

-1 -1
-2 -2
-3 -3
females -4
-5
-4
-5
eij – random error con-

% D % E
nected with the jth observa- 5
4
5
4
tion.
3 3
2 D=92, 12 2 D = 89,63
1 1
An F-test was applied 0

-1
0
-1
-2 -2
to test significance of the -3
-4
-3
-4
effect of the breeding lines -5 -5
on the analyzed features F G

5% 5%
(i.e. weight of eggs ex- 4
3 D=87,24
4
3 D=80,34
pressed in grams and as a

2 2
1 1
0 0
percentage of the female’s -1
-2
-1
-2
body weight, percentage -3

-4
-3
-4
-5 -5
of fertilization and percent-
age of live embryos after Figure 1. Constants of the least squares for the mass of eggs expressed in
grams (A) and as a percentage of the body weight of females (B) for the
24, 36, 48 and 56 hours of percentage of fertilization (C) and percentage of live embryos after 24 hrs
incubation). Duncan’s test (D), 36 hrs (E), 48 hrs (F) and 56 hrs (G) of the incubation of eggs from
modified by Kramer was three breeding lines (α - general mean).
applied to test significance of differences between means of the analyzed features as-
sessed for the three lines. The analysis enabled us to estimate the constants and means
of the least squares, which reflect the values of the traits within the analyzed main ef-
fect. Values of the means of least squares for the analyzed traits are given in Table 1,
253
Table 1. Least squares means (LSM) for mass of eggs (expressed in g and % of body weight of females) and for percentage of fertilization after 12 hours incubation and
254
percentage of live embryos after 24, 36, 48 and 56 hrs incubation of eggs (α-total means for the analyzed traits;; Se-least squares mean error).
Mass of eggs Percentage of fertilization Percentage of live embryos

Origin o females Mass of eggs (g)
(% b.w. of females) after 12 hrs fincubation
Breeding line α=812.31 after 24 hrs incubation after 36 hrs incubation after 48 hrs incubation after 56 hrs incubation
α=9.33 α=94.90
α=92.12 α=89.63 α=87.24 α=80.34
LSM Se LSM Se LSM Se LSM Se LSM Se LSM Se LSM Se
German line N 720.59 191.18 9.72 1.88 94.35 1.30 91.69 1.56 89.25 1.61 85.12 1.53 82.87 1.60
Hungarian line 7 709.16 226.74 7.15 2.62 96.42 1.81 92.43 2.17 90.42 2.24 89.35 2.13 75.82 2.24
Hungarian line 8 1007.16 181.82 11.12 1.79 93.93 1.24 92.25 1.48 89.22 1.53 87.24 1.45 82.31 1.52
Regression/body
42.27 102.22 -0.84 1.00 -2.01 0.69 -1.57 0.83 -1.83 0.86 -2.16 0.82 0.09 0.86
weight of females
Table 2. Correlations between the analyzed traits characterizing the effects of controlled reproduction of carp of the German line N, Hungarian line 7 and Hungarian line 8
Mass of Mass of Percentage Percentage of live embryos

Analyzed Breeding Mass of eggs of
traits line females (kg) (%b.w. of fertilization
eggs (g) females) after 12 hrs after 24 hrs after 36 hrs after 48 hrs after 56 hrs
incubation incubation incubation incubation incubation
1 2 3 4 5 6 7 8
N 0.24 -0.17 -0.29 -0.34 -0.42 -0.46 -0.67
1 7 -0.10 -0.37 -0.54 -0.35 -0.24 -0.44 0.65
8 0.24 -0.18 -0.76* -0.56 -0.66 -0.66 -0.22
N 0.65 -0.51 -0.36 -0.49 -0.50 -0.34
2 7 0.50 0.02 0.16 0.04 0.33 0.39
8 0.91* -0.18 0.17 0.17 0.02 -0.18
N -0.29 -0.33 -0.27 -0.18 0.01
3 7 -0.36 -0.30 -0.48 -0.29 -0.08
8 0.12 0.41 0.45 0.31 -0.08
N 0.96* 0.99* 0.88* 0.71
4 7 0.23 0.25 0.45 -0.36
8 0.87* 0.86* 0.82* 0.16
N 0.97* 0.85* 0.73
5 7 0.98* 0.95* -0.24
8 0.95* 0.90* 0.26
N 0.88* 0.75
6 7 0.92* -0.19
8 0.96* 0.41
N 0.94*
7 7 -0.14
8 0.46
whereas values of the constants of least squares are shown in Fig. 1. For each line, a
percentage of the fish which ovulated was calculated (Fig. 2), and phenotype correla-
tions between the analyzed features are contained in Table 2.
Table 3. Empirical values and predicted ones based on step regression equations for percentage of live embryos
after 56 hours of incubation R – multiple correlation coefficient .
Breeding lines
German N Hungarian 7 Hungarian 8
Resi- Resi- Resi-
Observed Predicted Observed Predicted Observed Predicted
dual dual dual
value value value value value value
value value value
(%) (%) (%) (%) (%) (%)
(%) (%) (%)
86 85,25 0,75 80 77,81 2,19 82 80,94 1,06
82 82,26 -0,26 70 69,65 0,35 80 83,02 -3,02
86 86,22 -0,22 80 79,72 0,28 79 81,72 -2,72
85 85,43 -0,43 80 76,62 3,38 84 81,00 3,00
81 85,76 -4,24 72 74,08 -2,08 84 85,86 -1,86
77 77,07 -0,07 74 78,13 -4,13 83 82,27 0,73
89 84,36 4,64
79 80,63 -1,63
80 80,18 -0,18
R 0,99 R 0,79 R 0,58
R2 0,98 R2 0,63 R2 0,34
In order to make predic- partial ov ulation (<200g )

tions on the percentage of complete ov ulation, bad quality spawn (f ertilization <25%)
live embryos after 56 hours complete ov ulation, good quality spawn
of ovulation within the ex- 100
amined lines, based on a set 90
of independent variables 80
such as a female’s body % 70
60
weight, weight of eggs (g), 50
40
percentage of fertilization 30
and percentage of live em- 20
10
bryos after 24, 36, and 48 0
hours of incubation, calcu-
lations were carried out by N 7 8
step regression. Entering Breeding lines
variables into an equation Figure 2. Percentage of females in which ovulation appeared after hor-
and removing them de- monal stimulation (data on very poor quality spawn have been excluded
pended on the critical val- from the calculations).
ues from the F-test. Solved equations were used to calculate predicted values, which
were then compared to the empirical ones for each observation (Tab. 3).
The values of the empirical data for such variables as spawn mass (g), percentage of
fertilization and percentage of live embryos after 56 hours of incubation as well as body
weight of females, weight of spawn (g) and percentage of live embryos after 56 hours of
255
Figure 3. Surface charts – for mass o eggs (g), percentage of fertilization and percentage
of live embryos after 56 hours of incubation – fitted by the square function.
a - line N: Z = -9803,5+0,86*x+203,80*y-7,5852E-6*x2-0,008*x*y-1,050*y2;
b - line 7: Z = -6458,3+0,54*x+139,15*y+2,99E-5*x2-0,006*x*y-0,7391*y2
c - line 8: Z = -339,57-0,23*x+11,59*y-1,82E-7*x2+0,0024*x*y-0,0742*y2
Figure 4. Surface charts – for the body weight of females, mass of eggs (g) and percentage
of live embryos after 56 hours of incubation – fitted by the square function.
a - line N: Z = -7361,1+317,65*x+128,26*y+0,62*x2-3,41*x*y-0,52*y2
b - line 7: Z = 3767,1-48,59*x-75,35*y-1,05*x2+0,80*x*y+0,36*y2
c - line 8: Z = -3737,5+471,75*x+38,92*y-8,46*x2-3,50*x*y-0,04*y2
256
incubation were a basis for deriving a square function and spatial prediction of the distri-
bution of points for these variables on a plane – separately for each line (Figs 3 and 4).
Results
Percentage of females from which spawn was obtained after stimulation
In the group of fish originating from line N, ovulation appeared in 80%, but partial
ovulation occurred in up to 20% of these females (Fig. 2). Partial ovulation was also
recorded in 10% of the females which belonged to line 7. In this line, the percentage of
fish in which complete ovulation occurred was 70% (Fig. 2). In the fish of line 8, 100% of
females dived eggs, and the weight of spawn from each female was satisfactory.
Times when complete ovulation appeared

Line N females produced spawn twice: four individuals produced spawn after 14
hours following the second injection of Ovopel, and two other ones – after another 4
hours (17 h). Likewise, in line 7 females, complete ovulation appeared twice. One fish
produced spawn 12 hours after the second injection of Ovopel, six other females – two
hours later. All the females originating from line 8 produced spawn 14 hours after the
releasing injection of Ovopel.
Effect of female origin on weight and quality of eggs

The results of the analysis of variance and F-test demonstrated that the effect of
the origin of the females was statistically significant (P≤0.05) on the weight of spawn
(expressed in grams as well as a percentage of the females’ body weight) and on the
percentage of live embryos after 48 and 56 hours of incubation of eggs (Tab. 4). The
values of the constant of least squares estimated for the examined lines indicate that the
highest weight of spawn (in grams and % of females’ b.w.) was obtained from the fish
which belonged to Hungarian line 8. The means of least squares for both traits defining
the weight of the spawn produced by this line were 1007.16g and 11.12% (Tab. 1) and
deviated in plus from the respective total means by 194.86g and 1.79% (Fig. 1 A, B).
Table 4. The F-test results.
Classification Analyzed traits

factor Mass of Percentage Percentage of live embryos
Mass of
eggs of after 48 hrs
eggs after 24 hrs after 36 hrs. of after 56 hrs
(% b.w. of fertilization incubation
(g)
females) after 12 hrs
Origin of
∗ ∗    ∗ ∗
females
Regression/
body weight   ∗  ∗ ∗ 
of females
*P≤0.05
The means of least squares for the weight of eggs produced by line N and line 7 fe-
males assumed similar values when the weight of eggs was expressed in grams (720.59 g
257
and 709.16 g, respectively, Tab. 1). When it was given as a percentage of the females’ body
weight, than a 2.57% higher value was determined for the fish originating from breeding
line N (Tab. 1). The average weight of eggs assessed for this trait for line 7 females deviated
in minus from the general average by 2.18% (Fig. 1B). The results obtained from Duncan’s
test, set in Table 5, demonstrated that the average mass of eggs expressed in grams ob-
tained from line 8 females differed significantly (P≤0.05) from the means calculated for the
other two lines. Statistically significant differences (P≤0.05) were also recorded between
the average mass of eggs expressed as % of the body weight of females originating from
line 7 versus the means calculated for lines N and 8 (Tab. 5).
Table 5. Results of Duncan’s test indicating significance of differences between the means for the analyzed traits
characterizing the effect of reproduction of three breeding lines of carp (*P≤0.05).
Breeding
Analyzed traits N 7 8
lines
• mass of eggs (g)  ∗
• mass of eggs (% b.w. of females) ∗ 
• percentage of fertilization  
• percentage of live embryos N
- after 24 hrs  
- after 36 hrs  
- after 48 hrs ∗ 
- after 56 hrs of incubation ∗ 
• mass of eggs (g) ∗
• mass of eggs (% b.w. of females) ∗
• percentage of fertilization 
• percentage of live embryos 7
- after 24 hrs 
- after 36 hrs 
- after 48 hrs 
- after 56 hrs of incubation ∗
The breeding lines did not determine significantly the percentage of fertilization or the
percentage of live embryos after 24 and after 36 hours of incubation (Tab. 4). It should be
emphasized, however, that the highest value of the mean fertilization percentage was re-
corded for the females of Hungarian line 7 (96.42%, Tab. 1). The means of the least squares
calculated for this trait in respect of the other two lines ranged around 94% (Tab. 1).
Approximately the same values of the average percentage of live embryos after 24
and 36 hours of incubation were recorded for all the three lines, with the mean value for
24 h incubation oscillating around 92% and for 36 h – around 90% (Tab. 1).
Statistically significant (P≤0.05) effect of the breeding line was demonstrated in the
case of the percentage of live embryos after 48 and 56 hours of incubation (Tab. 4). The
258
means of the least squares estimated for the percentage of live embryos after 48 hours
of incubation suggest that the highest mean value for this trait was determined for line
7, and the lowest one – for line N (89.35 and 85.12%, respectively; Tab. 1). The differ-
ence between these means was statistically significant (P≤0.05) (Tab. 5).
The lowest mean of the least squares (75.82%) computed for the percentage of live em-
bryos after 56 hours of incubation was determined for the females from line 7 (Tab. 1). It was
4.51% lower than the general mean (Fig. 1G). The other two means oscillated around 82%
(Tab. 1). The results of Duncan’s test showed that the mean assessed for this trait for line 7
differed significantly (P≤0.05) from the means recorded for lines N and 8 (Tab. 5).
Dependences between the analyzed features
Correlation coefficients between the body weight of the females and the weight of
eggs (expressed in grams and as a percentage of a female’s body weight) calculated for
the three lines can be considered as low (Tab. 2). The body weight of the females belong-
ing to all the three analyzed lines was negatively correlated with the five traits character-
izing the quality of spawn, except the positive correlation between the body weight of
females from line 7 and percentage of live embryos after 56 hours of incubation (Tab. 2).
The weight of eggs expressed in grams was positively correlated with all the other
analyzed traits but only in the group of fish originating from line 7 (Tab. 2).
The percentage of egg fertilization was positively correlated with the percentage of
live embryos after 24 hrs, percentage of live embryos after 36 hrs and percentage of live
embryos after 48 hrs of incubation of eggs for all the three lines. Statistically significant
(P≤0.05) correlation between the percentage of fertilization and each of the above traits
was noted for lines N and 8 (Tab. 2). The correlation between the percentage of fertilized
eggs and percentage of live embryos after 56 hrs incubation was not statistically signifi-
cant for any of the analyzed lines, but the highest positive correlation coefficient (0.71)
was found for the fish coming from German line N.
The percentage of live embryos after 24 hours incubation was positively correlated
with the percentage of live embryos after 36 hrs and the percentage of live embryos
after 48 hrs incubation within all the three analyzed lines. The correlation coefficients
between these traits for all the lines were very high (Tab. 2).
Likewise, the correlation between the percentage of live embryos after 36 hrs and
the percentage of live embryos after 48 hrs of incubation was statistically significant
(P≤0.05) for all the three breeding lines. The value of this correlation coefficient was the
lowest for line N (0.88) and the highest for line 8 (0.96) (Tab. 2).
Statistically significant correlation between the percentage of live embryos after 48 hrs
and percentage of live embryos after 56 hrs incubation was found only for line N (Tab. 2).
Regression predictions
During the calculations carried out by step regression in order to predict the percent-
age of live embryos after 56 hours of incubation, the following independent variable were
introduced into the regression equations: line N – body weight of females, percentage of
fertilization, percentage of live embryos after 48 hours of incubation; line 7 – body weight
of females, weight of eggs (g), line 8 – percentage of fertilization, percentage of live em-
259
bryos after 48 hours of incubation. The values of multiple correlation (R) and R2 for the
analyzed lines were, respectively: 0.99 and 0.98 for line N, 0.79 and 0.63 for line 7 and
0.58 and 0.34 for line 8 (Tab. 3). The results of the observed, predicted and residual values
are set in Table 3. Having compared these values, we could conclude that the most precise
prediction of the percentage of live embryos after 56 hours of spawn incubation was at-
tainable for the fish of line N, and the least precise one – for the fish of line 8.
Table 6. Main components of the square equation used for derivation of surface charts
(R – multiple correlation coefficient).
A – a set of variables: mass of eggs (g), percentage of fertilization, percentage
of live embryos after 56 hrs of eggs incubation
B – a set of variable: body weight of females, mass of eggs (g), percentage
of live ebryos after 56 hrs of eggs incubation
Arrangements Absolute
Breeding lines X Y R
of variables term
German N -9803.5 0.86 203.80 0.71
A Hungarian 7 -6458.3 0.54 139.15 0.52
Hungarian 8 -339.6 -0.23 11.59 0.22
German N -7361.1 317.65 128.26 0.86
B Hungarian 7 3767.1 -48.59 -75.35 0.64
Hungarian 8 -3737.5 471.75 38.92 0.22
Distribution of values of the analyzed traits in a spatial system

By analyzing the shape of spatially arranged planes, value of the absolute term, and val-
ues of variables X and Y, we could see that they were distinctly different for the analyzed lines
in both of the analyzed arrangements (Figs 3 and 4). This thesis can be confirmed by the val-
ues set in Tab. 6. The value of the multiple correlation enables us to state that the best fit of
the distribution of points to the plane is obtained for line N and the worst one – for line 8.
Discussion
Based on the results presented in this paper, it can be concluded that the highest
percentage (100%) of the fish in which complete ovulation appeared was among the
fish originating from Hungarian line 8, and the spawn from just one female of this line
had a low (<25) percentage of fertilization. It was only in this line, out of the three ex-
amined ones, that no partial ovulation was noticed in any of the females and all the fish
gave spawn at the same time. The results of other research which comprised 13 breed-
ing lines of carp, in which ovulation was stimulated with carp pituitary homogenate
(CPH), showed that Hungarian line 8 belonged to the lines in which the percentage of
produced spawn was low, not even reaching 50% (Brzuska 1991). A similarly low per-
centage of females of this line which produced spawn after stimulation with CPH was
observed in experiments carried out not only during (Brzuska 2004) but also outside the
reproductory season , i.e. in wintertime (Brzuska 1990). Noteworthy is the fact about
twice as many females of Hungarian line 8 started reproduction after being stimulated
with Lecirelin [D-Tle6, ProNHEt9] GnRH as among a group of fish which had undergone
hypophysation (64.5 and 37.5%, respectively) (Brzuska 2004).
In the experiment presented in this paper, complete ovulation among line 7 females
occurred in 70% of the animals. In another study carried out on 13 breeding lines of carp,
it was demonstrated that Hungarian line 7 was characterized by the highest (over 70%) per-
centage of females which produced spawn after hypophysation. Females of this line have
260
also been submitted to another study on induced ovulation during the summer season. The
results of this study proved that the percentage of spawned fish, which were subject to CPH
stimulation, reached nearly 70% (Brzuska and Adamek 1989). Analogously, in other experi-
ments on this line (where ovulation was induced with Ovopel) completed by Jamiński (2005)
and Cejko (2007), about 70% of ovulating females were observed. It is therefore noteworthy
that for a group of females from Hungarian line 7, included in an experiment performed
by Brzuska (2006a), the highest percentage (nearly 90%) of fish producing good quality of
spawn was obtained after stimulation with Ovopel, much higher than after hypophysation
or administration of Dagin [D-Arg6, Pro9Net-sGnRH + metoclopramide].
The breeding line N was a line within which 20% of females stimulated with Ovopel
had partial ovulation. Complete ovulation appeared in 60% of the fish, a result which
can be considered as satisfactory.
Our comparison of the average mass of eggs, in grams, which was produced by line
8 females with the average weights calculated for the other two lines demonstrated
that line 8 females produced the heaviest spawn (the difference between the means in
each comparison exceeded 280g). In another study, where ovulation was induced with
CPH and in which spawn obtained from females representing 13 lines was compared,
it was evidenced that the average weight of spawn for line 8 was about 120g heavier
than the average calculated for the whole set of fish. This line belonged to a group of
four lines which produced the highest weight of eggs (Brzuska 1991). The results of
another study on effects of propagation of line 8 carp after stimulating the ovulation
with CPH or Lecirelin showed that line 8 females stimulated with Lecirelin produced eggs
of a much higher weight (by ~300g) than the ones which underwent hypophysation
(Brzuska 2004). The females of this line used for out-of-season (winter) reproduction,
stimulated with CPH, produced much heavier spawn (>200g) than females of line X,
which were hybrids of a Polish and Hungarian line (Brzuska 1990).
The mass of eggs obtained from Hungarian line 7 females was the lowest among the
three examined lines. The results of a study completed on 13 breeding lines, CPH stimu-
lated, proved that this line belonged to the ones which produced an average weight
of spawn being about 160g higher than the total average mass (Brzuska 1991). High
weight of eggs produced by CPH-stimulated females of the same line 7 was also ob-
tained in a study carried out on the data collected during controlled reproduction in the
summer season (Brzuska and Adamek 1989). The results of the experiment carried out
on the same breeding line by Brzuska (2006a) demonstrated that a much higher weight
of eggs was obtained when females had received Ovopel than CPH or Dagin. Higher
spawn weight produced by line 7 females stimulated with Ovopel versus the ones receiv-
ing CPH was also demonstrated by Cejko (2007).
The average mass of eggs laid by the females from the German line N was similar to
the one produced by line 7 females only when weighed in grams. When converted into
the percentage of the body weight of females, the spawn mass of line N females was
much higher, reaching nearly 10% of the body weight of these females.
In respect of the quality of the spawn obtained in this experiment, it should be under-
lined that the mean percentage of fertilized eggs and live embryos after 24 and 36 hours of
incubation attained similar values for all the three lines. Significant differences in the quality
of spawn between the lines were noticed for such traits as the percentage of live embryos
after 48 and 56 hours of incubation. Considering the percentage of live embryos after 56 hrs
261
of incubation, it can be stated that the lowest average value for this trait was calculated for
line 7. The quality of spawn obtained from the same line after inducing ovulation with CPH
during the reproductive season was similar to the quality of spawn produced by three other
Hungarian lines, i.e. line T, line O and line 8 (Brzuska 1991). In a study completed by Jamiński
(2005) on four breeding lines originating from Gołysz, which were stimulated with Ovopel,
it was shown that the quality of spawn (expressed as % of live embryos after 24 hrs incuba-
tion) obtained from line 7 females was much poorer than the one produced by the French
line F and two Lithuanian lines, i.e. line B and line V. Some later research on Hungarian line
7 showed that the percentage of live embryos after 48 hrs of incubation was significantly
higher when ovulation was induced with Ovopel than with Dagin (Brzuska 2006a).
Noteworthy is the fact that the quality of spawn, expressed as a percentage of live
embryos after 56 hrs of incubation, which was demonstrated by line 8 females was high
and close to the quality of spawn laid by the females of the German line N. Satisfying
quality of spawn obtained from line 8 females stimulated with CPH had been signal-
ed before – both during (Brzuska 1991) and outside the reproductive season (Brzuska
1990). By analogy, a study carried out on the same line stimulated with CPH and Lecirelin
showed that an average percentage of live embryos after ovulation had been induced
with either of the preparations was higher than 80% (Brzuska 2004).
Recapitulating the above results, it can be claimed that the highest efficiency of
reproduction was shown by Hungarian line 8. Within this line, up to 100% females
underwent complete ovulation. The fish of this line also produced the heaviest spawn
(expressed in grams and as a percentage of the body weight of females), and the per-
centage of live embryos after 56 hrs of incubation was high. It is also important to notice
that the females originating from this line were the only ones which ovulated at the
same time. However, attention should be paid to the fact that for the same line, the least
precise prediction of the percentage of live embryos after 56 hrs of incubation was at-
tainable. The Hungarian line 7 was characterized by the poorest results of reproduction.
Although the percentage of females which underwent complete ovulation was satisfac-
tory, the average weight of spawn obtained from this line was the lowest and the quality
of spawn after 56 hrs of incubation was the poorest. The results of the reproduction of
the German line N (evaluated for the first time and thus not discussed before) can be
considered satisfactory although this line was the one where 20% of females had partial
ovulation. On the other hand, the prediction of the percentage of live embryos after 56
hours of incubation was the most precise for this line of carp.
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Białowąs H. 1999. Linie hodowlane w Polsce: linia węgierska 7. Przeg. Ryb. 1: 53-54.
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nus carpio L.) females in winter on the propagation results. Acta Hydrobiol. 32: 208-217.
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sertation No 153. 78 pp.
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Ovopelem; dalsze badania nad efektami rozrodu izraelskiej linii Dor-70 i jej krzyżówki z węgierską
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Brzuska E. 2003. Artificial propagation of the carp (Cyprinus carpio): two-year reproduction re-
sults of females of Hungarian line W and Polish line 6 after ovulation stimulation with carp pituitary
or mGnRH-a and dopaminergic inhibitor. Czech J. Animal Sci. 48: 139-151.
Brzuska E. 2004. Artificial spawning of carp (Cyprinus carpio L.); differences between the ef-
fects of reproduction in females of Hungarian, Polish and French origin treated with carp pituitary
homogenate or [D-Tle6,ProNHEt9]GnRH (Lecirelin). Aquacult. Res. 35: 1318-1327.
Brzuska E. 2006a. Artificial propagation of female Hungarian strain 7 carp (Cyprinus carpio) af-
ter treatment with carp pituitary homogenate, Ovopel or Dagin. Czech J. Animal Sci. 51: 132-141.
Brzuska E. 2006b. Badania efektywności stymulowanego hormonalnie rozrodu karpia (C. carpio
L.) różnego pochodzenia przeprowadzone w Zakładzie Ichtiobiologii i Gospodarki Rybackiej PAN w
Gołyszu w latach 1995-2005. In: Rozród, podchów i profilaktyka ryb karpiowatych i innych gatun-
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(Cyprinus carpio L.) oocytes with pregnant mare serum gonadotropin (PMSG). Acta Hydrobiol. 32:
437-446.
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Brzuska E., Grzywaczewski R. 1999. Artificial spawning of carp Cyprinus carpio L; differences
between the effects on reproduction in females of Israeli strain Dor-70 and its crossbred treated
with carp pituitary and Ovopel. Aquacult. Res. 30: 559-570.
Brzuska E., Białowąs H. 2002. Artificial spawning of carp, Cyprinus carpio (L.). Aquacul. Res.
33: 753-765.
Cejko B. 2007. Wpływ stymulacji hormonalnej samic różnych linii hodowlanych karpia (Cyprinus
carpio L.) na jakość ich ikry. PhD thesis, Faculty of Environmental Protection and Fisheries, University
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Horváth L., Szabó T., Burke J. 1997. Hatchery testing of GnRH analogue – containing pellets in
four cyprinid species. Pol. Arch. Hydrobiol. 44: 221-226.
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Instytut Genetyki i Hodowli Zwierząt PAN, Jastrzębiec. 64 pp..
Irnazarow I., Białowąs H. 1995. Genetic characteristic of carp breeding lines at the Institute of
Ichthyobiology and Aquaculture of the Polish Academy of Sciences Gołysz. 2. Hungarian lines. Acta
Hydrobiol. 37: 141-151.
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lanego (Cyprinus carpio L.). Master thesis. Akademia Rolnicza im. H. Kołłątaja w Krakowie. Wydział
Hodowli i Biologii Zwierząt. 54 pp.
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(Cyprinus carpio L.) to Trypanoplasma borreli: Influence of transferrin polymorphisms. Vet. Immu-
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2006. Genetic differences in natural antibody levels in common carp (Cyprinus carpio L.). Fish &
Shellfish Immunology 21: 404-413.
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263
Optimization of the reproduction
of chub (Leuciscus cephalus L.)
under controlled conditions
Sławomir Krejszeff1, Dariusz Kucharczyk1,
Katarzyna Targońska1, Daniel Żarski1,
Krzysztof Kupren1, Andrzej Mamcarz1,
Sebastian Piszczała2
1
Protection and Fisheries, University of Warmia and Mazury in Olsztyn,
Knieja Fish Farm
2
Abstract
The objective of this study has been to optimize reproduction of chub under controlled
conditions through administration of one of the three different hormonal stimulants: carp
pituitary homogenate CPE, human chorionic gonadotropin hCG and the mammalian ana-
logue of gonadotropin-releasing hormone in a complex preparation with metoclopramide
(Ovopel), and through the application of reproductive procedures in different time during
the spring season (from March to June). The experiment was run for three subsequent
reproductive seasons on fish originating from 8 different populations. The results point to
the efficacy of CPE and Ovopel, although statistically better effects were obtained when
reproduction was stimulated with Ovopel, as the percentage of survivability of embryos
was than much higher (on average 87%) compared to CPE (on average 79%). However,
the latency time (from the releasing injection) was much shorter when CPE rather than
Ovopel was administered (16-18h versus 24-32h, respectively). Reproduction stimulat-
ed before the proper reproductive season (from March to May) led to high mortality of
spawners (over 75%, both among females and males) and a very low effectiveness of the
reproductive procedures (the percentage of ovulations and survivability among embryos
below 10%). The best results were achieved when reproduction was run in June (61% of
ovulations and 95% survivability of embryos). The results reported in this paper can con-
siderably enhance the effectiveness of chub reproduction under controlled conditions.
Introduction
The future fate of many fish populations living in the wild is uncertain. In just five
years (1992-1997), the number of freshwater species considered as not threatened in Po-
land decreased from 23 to 18 (Błachuta 1998). Developing reproduction biotechnologies
265
with the aim of acquiring possibly the best quality gametes is the base for fish restitution
programmes (Kucharczyk 2002). For many years, it has not been easy to find in the Polish
and world literature papers dealing with the artificial reproduction of non-commercial fish
species inhabiting our country’s open waters. The fear that some of these species may
disappear has stimulated more interest in this field. Nevertheless, advanced work on the
reproduction biotechnology has been attained in the case of just a few species, including
ide, asp, dace and nase (Targońska-Dietrich et al.2004, Cejko et al.2008, Jamróz et al.2008,
Krejszeff et al.2008, Kucharczyk et al.2008, Targońska et al.2008, Żarski et al.2008, 2009).
Chub is probably the next species whose reproduction will soon be elucidated well enough
to make rational management of its populations feasible.
In a hatchery, fish reproduction is possible mainly through hormonal stimulation (Ku-
charczyk 2002). Among the earliest preparations applied to induce ovulation in females
and spermiation of males was pituitary homogenate. At present, there are many avail-
able hormonal preparations which can successfully replace pituitary extracts (Kucharczyk
and Szabo 1998, Brzuska and Adamek 1999, Brzuska 2000).
Stimulation of sexual maturation can be attained by affecting the hypothalamus, pitui-
tary or gonads. The hypothalamus is most often stimulated by changes in the environmen-
tal conditions, mainly temperature and photoperiod. Hormonal preparations, in turn, are
administered when the stimulation is to be achieved on the other two levels: the pituitary
– with natural or artificial gonadotropin-releasing hormones, and gonads – with fish or hu-
man gonadotropins (Bieniarz and Epler 1991). Part of the development of a reproduction
biotechnology involves testing how these different hormonal stimulants work. Doses and
type of hormones are determined according to a fish species and its reproductive strategy.

Acquisition of spawners
During our three-year study, chub spawners were obtained from the rivers Wałsza,
Vistula, Rozpuda, Marózka, Łyna, Narew and Grabia. All the fish were transported to
a hatchery at the Chair of Lake and River Fisheries, where they were placed in tanks hold-
ing 1,000 dm3 of water. Each tank was equipped with devices for aeration, temperature
(± 0.1°C) and photoperiod regulation (Kujawa et al. 1999).
Fish manipulations
Each time, prior to administration of hormonal stimulants, the oocyte maturity stage
was assessed on a four-stage scale (Brzuska and Bieniarz 1977):
Stage 1 – the nucleus placed centrally
Stage 2 – the nucleus slightly shifted (between the centre and middle of the radius)
Stage 3 – the nucleus behind the middle of the radius
Stage 4 – the nucleus on the edge or is absent (has undergone degradation)
Oocytes were sampled with a catheter, according to the methodology described by
Kujawa and Kucharczyk (1996). Samples of egg cells were taken in vivo and placed in
Serra liquid in order to clarify the cytoplasm. After 5-minute exposure, the position of
the nucleus was assessed on the scale presented above. Females whose oocytes were
on 2-3/3 stage of maturation (the best moment for application of hormonal stimulation
266
Optimization of the reproduction of chub (Leuciscus cephalus L.) under controlled conditions
in cyprinid fish; Kozłowski 1994) were given hormonal injections. Before the manipula-
tions, the fish were anaesthetized in a 2-phenoxyethanole solution (0.5cm3/dm-3 dose).
Hormonal stimulation
Two variants of chub reproduction were tested. In the first one, conducted during the
natural spawning season of that fish, different hormonal preparations were administered:
carp pituitary homogenate (CPE), human chorionic gonadotropin (hCG), a GnRH analogue
with metoclopramide (Ovopel) (Tab. 1). Once they reached the appropriate stage of gonad
maturity, the spawners were divided into four groups and subjected to hormonal stimula-
tion. In the second variant of the experiment, an attempt was undertaken to determine
the best moment for artificial reproduction. For this purpose, reproduction was run in
monthly intervals from March to June. Chub spawners were brought to the hatchery every
few weeks from February to early June. The fish were maintained in tanks, raising the
water temperature and extending the photoperiod. When the oocytes reached the matu-
ration stage 2 or 3, the fish
were given Ovopel. Stimu-
lation was run according
to the doses presented in
Table 2. The injections were
given intraperitoneally un-
der the vertal fin (Fig. 1). All
the hormonal preparations
were homogenized and dis-
Figure 1. Making a hormonal injection of Ovopel suspension to a chub
solved in sterile 0.9% NaCl. female (Photo: the authors).
Table 1. Hormonal preparations used for controlled reproduction of chub Leuciscus cephalus (L.).*
Initial injection Stimulating injection
Hormonal agents 0 hour 24 hours
females males females males
CPH (mg kg-1) 0.4 - 3.6 2.0
HCG (IU kg ) -1
200 - 1800 1000
Ovopel (pellet kg-1) 1

/10 pellet - 1 pellet 1
/2 pellet
* the control group fish were given injections of 0.9% NaCl.
Table 2. Doses of Ovopel (pellet kg-1) administered in reproduction of chub Leuciscus cephalus (L.).
Initial injection Resolving injection
0 hour 24 hours
females males females males

1
/10 - 1 /2
1
267
Production of gametes
Semen was sampled with syringes calibrated to 0.1 ml. Before checking its motility,
semen was stored in a refrigerator at about 4oC. Motility of spermatozoa was determined
under a microscope (500x magnification), by diluting the semen at a 1 : 1 ratio in 0.5%
NaCl solution immediately
before an observation. By
pressing the abdominal
walls of fish Spawn was
obtained and transferred
into dry plastic contain-
ers (Fig.2), in which it was
stored until fertilization.
Fertilization was performed
with the dry method.
Incubation
Samples of eggs (about
100-150 grains), from each
female, were placed on
Petri plates in two repli-
cates. Afterwards, 0.05 cm3
of a mixture of semen and
water was added in order Figure 2. Stripping of eggs from the female of chub previously
to activate the gametes. stimulated with Ovopel (Photo authors).
When water is added, it reacts with the glycoprotein on the surface oocytes so that chub
spawn becomes sticky and incubation on Petri plates is possible. Marked samples were
placed in a tank with a constant supply of fresh water. The incubation temperature was
16oC. After the eyed-egg stage was reached (pigment appeared in the eyes of embryos),
the percentage of live embryos was counted.
Statistics
Statistical differences between the groups (survivability of embryos to the eyed-egg
stage) was analyzed with Duncan’s test at the level of significance below 5% (p<0.05).
Results
The differences in the efficacy of the applied stimulants, observed in this study, are
shown in Table 3. When hCG was administered, a small increase in the percentage of
males producing milt was observed versus the control. The results were 68±6% (hCG)
and 53±6% (control). By comparison, administration of carp pituitary extract or Ovopel
caused spermiation in all males. The same regularity was attested in respect of the
amount of produced semen. In the group injected hCG, the amounts of obtained semen
were similar to those produced by the control group (2.5±0.4 and 2.4±0.7 cm3 kg-1 body
weight of spawners). Significant differences, with regard to the control, were observed
in the other two groups. The amount of semen from the males administered carp pitui-
tary extract was 3.7±0.3 cm3 kg-1, and from the ones injected Ovopel – 3.6±0.2 cm3 kg-1.
268
The lowest motility of spermatozoa was recorded in the control group (6-70%). In the
other groups, it was much higher (80-90%).
With respect to females, no germ products were obtained from the group which
spawned spontaneously and the one stimulated with hCG. Positive effect, however, was
achieved in the other two groups. The most effective hormonal stimulant proved to be
Ovopel (36% ovulating females). Significant differences were observed in terms of the
applied stimulant and the survivability of embryos until the eyed-egg stage. In the group
of fish injected pituitary extract, 79±3% of embryos were viable during that stage, as
compared to 87±4% of embryos produced by fish stimulated with Ovopel. Differences
were also noticed in the time when ovulation occurred depending on the applied hor-
monal stimulation (Tab. 3). Females which were given pituitary extract ovulated within
16-18h, while the ones stimulated with Ovopel – within 24-36h.
The results of the second variant of the experiment are presented in Table 4. The
percentage of males releasing semen was the lowest in March (68%) and April (74%).
In contrast, all males produced semen in May and June. The amount of obtained semen
rose gradually every month. In March, it equalled just 2±0.3 ml kg-1, but in June it rose
to 3.1±0.3 ml kg-1. Motility of spermatozoa was the lowest in March and April (70 and
70-80%, respectively), and the highest in May and June (80-90%). The percentage of
ovulating females was very low in April and May (just 8%), and the amount of obtained
spawn was small. Many more females ovulated in June (61%), and oocytes in non-ovu-
lating females were seen to be maturing. The survivability of embryos to the eyed-egg
stage was low in April and May (3.6 and 8.9%, respectively) but much higher in June
(95%). High mortality among spawners was recorded during the whole study (37-53%
of males and 53-68% of females).
Table 3. Results (mean ± SD) of a three-year-long study into the controlled reproduction
of chub Leuciscus cephalus (L.) during the natural reproductive season.
Group of fish Control CPE HCG Ovopel
Percentage of spermiating males (± SD) 53 ± 6 100 68 ± 6 100
Amount of semen (ml kg ) (± SD)

-1
2.4 ± 0.7 b
3.7 ± 0.3 a
2.5 ± 0.4 b
3.6 ± 0.2a
Motility of spermatozoa % (± SD) 60-70 80-90 80-90 80-90
Mortality of males (%)* 37 42 36 53
Percentage of ovulations 0 28 0 36
Maturation of oocytes little or
yes little yes
in non-ovulating females none
Time lapse between injection
- 16-18 - 24-32
and ovulation (h)
Survivability of embryos to eyed-egg stage
- 79 ± 3b - 87 ± 4a
(%) (± SD)
Mortality of females (%)* 53 47 49 68
Data marked with the same letter do not differ statistically (P < 0.05)
* mortality mainly while fish are kept after spawning, due to body damage
269
Table 4. Results (mean ± SD) of controlled reproduction of chub Leuciscus cephalus (L.) in different months.
Reproduction March April May June

Percentage of spermiating males 68 74 100 100
Amount of semen (ml kg ) (± SD)
-1
1.2 ± 0.3 1.6 ± 0.4 2.9 ± 0.2 3.1 ± 0.3
Motility of spermatozoa % 70 70-80 80-90 80-90
Mortality of males (%)* 76 86 76 48
full or par-
Type of ovulation none partial partial
tial
Percentage of ovulations - 8 8 61
Maturation of oocytes
yes yes yes yes
in non-ovulating females
Survivability of embryos
- 3.6 8.9 95
to eyed-egg stage (%)
Mortality of females (%) *
85 76 100 11
* mortality mainly while fish are kept after spawning, due to body damage
Discussion
The results of our study on chub reproduction coincide with other reports on repro-
duction of cyprinid fishes, which indicate that after hormonal stimulation males produce
more ejaculate, frequently of a superior biological value (Kozłowski 1994, Kucharczyk et
al. 1997a, Kucharczyk 2002, Cejko et al. 2008). Similar observations have been reported
in the case of chub. Different hormonal preparations stimulate spermiation to a various
degree. The weakest effect was produced by hCG, whereas the results generated by CPE
and Ovopel did not differ much.
The results of hormonal stimulation were even more varied in respect of females. The
control group females did not commence reproduction. Meanwhile, although the fish
were kept in suitable environmental conditions, the nucleus in female germ cells was
not seen to have changed its position. The hCG stimulated females responded similarly.
Although hCG was noticed to have some effect on the maturation of female gametes,
such as a shift of the nucleus towards the peripheral area, the stimulant was not suf-
ficiently effective and did not trigger ovulation. A similar situation was observed among
bream females (Kucharczyk et al. 1997a, 2005). The efficacy of this stimulant, however,
has been demonstrated on cyprinid fish with batch spawning, i.e. crucian carp, tench
and rudd (Kucharczyk et al. 1997b, 2007, Kucharczyk 2002, Targońska et al. 2009).
Application of fish pituitary extracts has been known and used for many years. How-
ever, apart from having some advantages, this method has several drawbacks. By intro-
ducing hormones to fish organisms, we also introduce a whole range of proteins and
other compounds which the pituitary contained. This may cause allergies. Each subse-
quent administration of pituitary extract can be less tolerable to fish due to an increasing
level of antibodies. Another disadvantage is that after hCG is administered, oocytes in
ovaries undergo hydration, whether or not the fish will ovulate. If germ cells are released
when no ovulation occurs, the fish can experience different health problems and may
not be able to reproduce in the following spawning seasons. In some cases, they can
even die (Kucharczyk 2002). CPH produces a direct effect on gametes and causes their
270
maturation. This effect has been termed as direct influence. Pure, isolated hormones
(LH or FSH) directly affecting gametes produce a different effect than their analogues
(Kucharczyk et al. 1998, Yaron et al. 2009, Krejszeff et al. 2009). By affecting endocrine
glands, they stimulate them to produce own hormones, which in turn stimulate matura-
tion of gametes. This is the reason why fish administered CPH reach spawning maturity
sooner than fish stimulated with GnRH or its analogues. This difference is distinctly seen
when the time lapse between an injection of a stimulant and production of spawn is
measured. In all the cases described in this article, ovulation of fish stimulated with
Ovopel appeared a few hours later than that of the fish injected CPH. The same depend-
ence has been noticed not only for cyprinid fish but also in the case of perch (Kucharczyk
et al.1996, 2005, 2008, Jamróz et al. 2008, Targońska et al. 2008, Żarski et al.2008,
2009). As a rule, the efficacy of both groups of hormonal stimulants (CPE and Ovopel)
was similar among chub males. Nonetheless, Ovopel proved to be more successful. More
males were ready to spawn when administered Ovopel, and the biological quality of
spawn, understood as the survivability of eyed-egg embryos was much better.
High mortality of spawners was observed during this study. In fact, it was higher
than found among other species of fish (Kucharczyk 2002). As our observations proved,
in some groups, the mortality of spawners was as high as 100%.Some cases of death
could have been caused by damage, both external and internal, that the fish suffered
when they were caught from open waters. A small percentage of females which begin
to spawn as well as small quantities of spawn obtained from most of the fish (partial
ovulation) compounded by the high mortality of spawners limit the success of chub re-
production under controlled conditions. Therefore, more research on this species should
be done to test hormonal preparations, ways of obtaining fish for spawning and keep-
ing spawners in tanks.
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272
Hybridization among fish and its
importance to biodiversity
Lucyna Kirczuk, Józef Domagała
University of Szczecin. Chair of General Zoology, ul. Felczaka 3c,

71-412 Szczecin, tel. +48 91 444-16-22; fax +48 91 444-16-23;
e-mail: lucyna.kirczuk@univ.szczecin.pl
Abstract
Because of their life habitats and external fertilization, hybridization among fish is
much more common than among other vertebrates. Identification of hybrids relies on
analysis of coloration, conformation, calculable and measurable traits as well as karyo-
logical, biochemical and molecular analyses.. Hybrids represent different types of fertil-
ity. The most common are sterile individuals. Some hybrids are fertile or else they have
unequally developed gonads. Fertile hybrids in the natural environment are a threat to
pure species and biodiversity due to possible introgression.
Introduction
Crossing of different fish species in nature is a common but undesirable event (Camp-
ton 1987, Elo et al. 1995). Unfortunately, human activity largely contributes to an in-
crease in the hybridization level. Destroying proper water relations, changing the height
of the water table in reservoirs and thus limiting fish’s breeding grounds as well as shift-
ing the borders of an area where a certain species dwells, thus causing an overlap with
an area occupied by another species raises the degree of hybridization (Garcia de Léaniz
and Verspoor 1989; Hammar et al. 1991). Introduction of new fish species is especially
dangerous to native ones because of the risk of hybridization (Dowling and Secor 1997;
Janowicz 2004). Sympatric species, whose geographical areas of occurrence overlap, are
at the highest risk (Hammar et al. 1991, Delling et al. 2000). Hybridization means that
hybrids are created, which in itself is a threat to pure species and biodiversity (Janowicz
2004). On the other hand, crossing is used in biotechnology in order to produce new in-
dividuals with expected traits. Therefore, when making decisions on a local level regard-
ing hydrotechnical projects, fish catches, introduction of new species, etc, it is important
to remember how such decisions may affect biodiversity. For this purpose, it is necessary
to know how to identify hybrids. In turn, our knowledge of their reproduction capacity,
by analyzing the development of their gonads, enables us to create back-crossbreeds
and to preserve the biodiversity of a given ecosystem.
273
Methods for identification of hybrids

In the natural environment, hybrids are often difficult to distinguish, for example
among cichlids in lakes in East Africa (Eccles and Trewavas 1989). Therefore, different
ways of identification of hybrids are used. Among the earliest criteria for identification
of hybrids was morphology, based on which Alm (1955), as one of the first researchers,
described salmonid hybrids originating from natural habitats. In most of the studies on
hybrids carried out in later years, fish’s external appearance was also analyzed, includ-
ing coloration (Hedenskog et al. 1997), shape of the body (Chevassus 1983; Wilkins
et al. 1994), the morphology of processes on the branchial arch (Kazakov et al. 1984),
measurable traits (Wilkins et al. 1994; Hedenskog et al. 1997) and calculable ones, which
are particularly useful when the parental species are different from each other in this
respect and the characteristics of the hybrids are intermediate between the parental
forms (Leary et al. 1983; Ma and Yamazaki 1986; Nelson 1990; Delling et al. 2000).
Apart from morphological characteristics, investigations dealing with hybrids rely on the
skull’s osteologic bones (Penczak 1978; Dorofeyeva et al. 1989, 1990) or on the level
of variability in the size of ovulating spawn (Kazakov i Ilyenkova 1982). The physiologi-
cal criterion for distinguishing hybrids is based on the evaluation of the duration of the
embryonic development, which – according to Makeevej (1975) - in the hybrid Cyprinus
carpio x Hypophtalmichthys molitrix is in between those of the parental species. In Po-
land, in habitats shared by rudd and bream, the fact that they can cross and produce
hybrids is suggested by a different type of spawning or differences in vitellogenesis
(Kopiejewska et al. 2007).
Identification of hybrids is also performed via examination of karyotypes, including
analysis of the number of chromosomes and their morphology (Johnson and Wright
1986; Hartley 1996) or through biochemical assays based on the electrophoretic de-
termination of the motility of particular enzymes (Nelson 1990; Hammar et al. 1991;
Delling et al. 2000). This method is very useful provided that the parental species are
polymorphic in respect of the motility of these enzymes because the hybrids inherit them
from both parents (Chevassus 1983). Hybrids have also been identified on the basis of
the assessment of the diameter of nuclei in erythrocytes, which in Cyprinidae hybrids
had intermediate values, or according to the cytometric analysis of DNA in the cellular
nucleus, which can help us to determine the ploidy of hybrids (Allen 1983). Much infor-
mation on the dynamics of inter-species interactions and the direction of hybridization
is provided by analysis of mitochondrial DNA (mtDNA) in hybrids, because it is inherited
only from the mother (Nelson 1990; Jansson i Öst 1997). At present, a commonly ap-
plied tool for identification of hybrids is analysis of nuclear DNA, and especially micros-
atellite markers.
Fertility of hybrids
Fertility of fish hybrids is an extremely interesting question. In respect of their repro-
duction capacity, fish crossbreeds can be divided into several groups. The first group,
and the most numerous one, consists of sterile individuals. It comprises hybrids with
atrophic gonads containing germ cells halted during the early stages of gametogen-
esis; this is known as gonadal sterility (Chevassus 1983). Sterile hybrids encompass such
crossbreeds as Pseudogobio esocinus x Gnathopodon elongatus (Suzuki 1976), Barbus
longiceps x Capoeta damascini (Stoumboudi et al. 1992), Oryzias latipes x O. curvinotus
274
Hybridization among fish and its importance to biodiversity
(Hamaguchi i Sakaizumi 1992), di- and triploid hybrids Sparus aurata x Pagrus major
(Gorshkov et al. 2002) or Pleuronectes ferrugienus x Pleuronectes americanus (Park et
al. 2003). Sterile crossbreeds are frequent among Salmonidae, e.g. Salmo trutta x Sal-
velinus fontinalis (Blanc and Chevassus 1986) or Oncorhynchus masou x O. gorbuscha
(Ma and Yamazaki 1986). Some other crossbreeds are sterile due to an incomplete sexual
development (only some cells mature) (Hubbs 1955). This type of sterility appears in
the hybrids of Centrarchidae (Capanna et al. 1971). Zygotic sterility is represented by
the crossbreeds Ctenopharyngodon idella x Cyprinus carpio (Hubbs 1933) as well as
Alburnus alburnus x Rutilus rubilio (Berrebi et al. 1989), in which, despite well-devel-
oped gonads and viable gametes, the zygotes were not viable. Noteworthy is the fact
that individuals of the same cross and at the same age, may have differently developed
gonads, e.g. some males of Salmo trutta x Salvelinus fontinalis underwent spermiation
while others had thread-like testes (Suzuki and Fukuda 1973), or else, had testes which
were underdeveloped, partly developed or produced watery sperm, like the hybrids On-
corhynchus mykiss x Salvelinus alpinus (Blanc and Chevassus 1982). Such differentiation
has also been detected in females, for example the hybrids Oncorhynchus masou x O.
gorbuscha, which comprised both sterile and ovulating individuals (Ma and Yamazaki
1986). Among the hybrids present in Poland, crossbreeds of Rutilus rutilus x Leuciscus
idus likewise underwent an uneven sexual development, i.e. germ cells of some females
matured only in a fragment of a gonad whereas males reached the stage of spermiation
and spermatocytes (Kopiejewska et al. 2004a). In the case of hybrids Scardinius eryth-
rophthalmus x Abramis brama, studies have demonstrated that they can reach sexual
maturity in the natural environment. Reaching sexual maturity by hybrids also depends
on the gender, e.g. only few males of reciprocal hybrids Salvelinus fontinalis x Salvelinus
alpinus underwent spermiation, in contrast to females, most of which were not fully
sexually mature (Dumas et al. 1996).
Evaluation of the fertility, quality of gametes and possibilities of obtaining F2 genera-
tions or backcrosses among hybrids is of particular importance in aquaculture (Suzuki
and Fukuda 1973; Chevassus 1979; Blanc and Chevassus 1982). During the backcross-
ing of hybrids of Cyprinidae, a 5-10% fertilization rate was achieved (Bakos et al. 1978)
but the development did not surpass the stage of forming the blastodisc. By analogy
to other species, this could be the consequence of differences in the karyotypes of the
parental species and difficulties in coupling chromosomes during meiosis (Nygren et al.
1975; Johnson and Wright 1986). Examples of fertile hybrids with gonads and gametes
of normal structure and size, capable of producing F2 offspring and backcross are native
hybrids Coregonus lavaretus x Coregonus peled (Falkowski et al. 1995). As Hammar et
al. (1991) claim, the offspring of inter-species hybrids has depressed viability, fertility
and fitness due to a new combination of loci, which also influences phenotype varia-
tion. Poorer survivability relative to the parental forms has been determined for hybrids
Sparus aurata x Pagrus major (Gorshkov et al. 2002). According to Senan et al. (2004),
inferior fitness of hybrids macrocephalus x Clarias gariepinus prevents introgression.
There are, however, crossbreeds like Coregonus lavaretus x Coregonus peled (Falkowski
et al. 1988), which demonstrate comparable or superior survivability compared to their
parental species.
Finally, there are also hybrids such as Morone saxatilis x Morone chrysops, which
demonstrate favourable traits (Chervinski et al. 1989). In biotechnology, it is important
275
that the hybrids can acquire traits which fish breeders will find valuable because hybridi-
zation has become one of the biotechnological methods. Owing to hybridization, popu-
lations of fish have been obtained which have improved tolerance to chill stress, salinity
(Ma and Yamazaki 1986), concentration of ammonium (Hargreaves and Kucuk 2001) or
which have a better growth rate and higher tolerance to diseases (Hulata 2001). On the
other hand, presence of hybrids in the natural environment may be associated with the
risk of supplanting pure species and introgression, which can be a threat to biodiversity
in a given site (Janowicz 2004). It should be taken into account that higher incidence of
hybrids is very often connected with hydrotechnical changes, introduction of new spe-
cies and inappropriately conducted restitution (Behnke 1968; Makeeva 1975; Stanley
and Jones 1976; Hammar et al. 1991; Kazakov 1998). Conservation of biodiversity and
habitats is based on a broadly understood planning process concerning hydrotechnical
projects as well as the evaluation of the effect of introduced fish and hybrids, even the
ones that possess favourable characteristics, on indigenous fish species. Hybridization
among fish living in the natural environment is an unwanted event because it diminishes
populations of pure species and threatens biodiversity.
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Contact
authors e-mail
Leszek Augustyn leszek.augustyn@pzwns.info.pl
Magdalena Bowszys mbowszys@uwm.edu.pl
Elżbieta Brzuska urszula.karosek@fish.edu.pl
Beata Irena Cejko b.cejko@pan.olsztyn.pl
Przemysław Czerniejewski pczerniejewski@zut.edu.pl
Krystyna Demska-Zakęś krysiadz@uwm.edu.pl
Józef Domagała troc@univ.szczecin.pl
Katarzyna Dziewulska katarzyna.dziewulska@univ.szczecin.pl
Grażyna Furgała-Selezniow graszka@uwm.edu.pl
Jan Glogowski j.glogowski@pan.olsztyn.pl
Paulina Hofsoe paulina.hofsoe@zut.edu.pl
Jolanta Kempter jolanta.kempter@zut.edu.pl
Sławomir Keszka skeszka@zut.edu.pl
Lucyna Kirczuk lucyna.kirczuk@univ.szczecin.pl
Radosław Kajetan Kowalski r.kowalski@pan.olsztyn.pl
Sławomir Krejszeff darekk@uwm.edu.pl
Jacek Kubiak jacek.kubiak@zut.edu.pl
Dariusz Kucharczyk darekk@uwm.edu.pl
Roman Kujawa reofish@uwm.edu.pl
Krzysztof Kupren krzysztof.kupren@uwm.edu.pl
Sylwia Machula sylwia.machula@zut.edu.pl
Andrzej Mamcarz mamcarz@uwm.edu.pl
Włodzimierz Marszelewski marszel@umk.pl
Andrzej Martyniak kbhr@uwm.edu.pl
Maciej Mickiewicz mickiewicz@infish.com.pl
Katarzyna Mierzejewska katarzyna.mierzejewska@uwm.edu.pl
Bartosz Mysłowski bartosz.myslowski@zut.edu.pl
Arkadiusz Nędzarek arkadiusz.nedzarek@zut.edu.pl
Katarzyna Palińska katarzyna.palinska@uwm.edu.pl
Rafał Pender rafal@pzwszczecin.com
Przemysław Pol reofish@uwm.edu.pl
Mirosław Półgęsek miroslaw.polgesek@zut.edu.pl
Jacek Sadowski jacek.sadowski@zut.edu.pl
Beata Sarosiek b.sarosiek@pan.olsztyn.pl
Jerzy Sell sell@biotech.ug.edu.pl
Andrzej Skrzypczak sandacz@uwm.edu.pl
Adam Solarczyk adamsolarczyk@poczta.onet.pl
Robert Stabiński pzw@suwalki.com.pl
Katarzyna Stańczak kbhr@uwm.edu.pl
Mirosław Szczepkowski szczepkowski@infish.com.pl
Mariusz Szmyt mariusz.szmyt@uwm.edu.pl
Renata Tandyrak renata.tandyrak@uwm.edu.pl
Adam Tański atanski@zut.edu.pl
Katarzyna Targońska targonska.k@uwm.edu.pl
Agnieszka Tórz Agnieszka.Torz@zut.edu.pl
Konrad Turkowski kontur@uwm.edu.pl
Arkadiusz Wołos awolos@infish.com.pl
Bogdan Wziątek b.wziatek@uwm.edu.pl
Zdzisław Zakęś zakes@infish.com.pl
Daniel Żarski danielzarski@interia.pl
Project no PL0468 „Ichthyological biodiversity of lakes - elaborating a model for problem solution:
a case study of natural resources of autochtonous common whitefish in Lebsko Lake (Lebsko Lake
whitefish) (acronim Fish-WILL), under the EEA Financial Mechanism and Norway Grants, priority
sector „Environment and sustainable development”
279

Fish Management in A Variable Water Envi

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Fish Management in A Variable Water Envi

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Fish management

Translating: mgr Jolanta Idźkowska

© 2011 Faculty of Environmental Protection and Fisheries

Department of Lake and River Fisheries,

Biodiversity of lake ichthyofauna – different scales of the problem

Group Proportion threatened (%)

al. 2003). In the 1970s there

Various changes in the

Family Species Temperature preference

Family Species Threat Legal status

Effect of the management of open waters on biodiversity of ichthyofauna

sources (12th-13th centuries)

Table 5. Taxonomic account of both exploited and protected ichthyofauna

Family Species Threat Legal status

Family Species Year (Period)

Fish stock releases

Table 7. Taxonomic account of ichthyofauna stocked to Mazurian fresh waters

Family Species 1964 - a 1975 After 1989

Where we are heading – or reaching a compromise

of biodiversity is more and more often analyzed in terms of economic transformations

Species Reason for introduction Impact

Ordinance 1997. Rozporządzenie Ministra Rolnictwa i Gospodarki Żywnościowej z dnia 10 grud-

University of Gdańsk, Department of Genetics,

Effects of fish stocking on the genetic structure of a population

in the abundance of a population between subsequent generations, no overlap of the

Applicability of microsatellite DNA as a genetic marker to conservation of ich-

The structure of microsatellite DNA

Advantages of microsatellite DNA as a genetic marker

Evaluation of the genetic variation of a population

Natomiast mtDNA jako marker genetyczny cechuje się mniejszą informatywnością

Evaluation of the long-term changes in the genetic structure of populations

Evaluation of modifications in the genetic structure of populations going backwards

Biodiversity of hatchery production

The technical infrastructure of hatcheries versus hatchery production structure

It should also be no-

West Pomeranian University of Technology in Szczecin,

The fish stocking management in the context of biodiversity

Local populations and their genetic differentiation

which enables its identification is a consequence of the morphological or geographical

Abundance of a population versus its being threatened

individuals, and reproduction based on 1 or 2 females and a few males is dangerous. As

Molecular markers used in studies on rare and protected species

Effect of fishery management on genes, species and ecosystems

• Mitigation (to compensate for losses in populations caused by environmental changes);

Materials and methods

Number of enterprises Combined surface area of lakes exploited

Number of enterprises Combined surface area of lakes exploited

Results and discussion

During the 2001-2009 period, fisher-

Table 3. Types of stocking material (X) of particular fish species released

Mickiewicz M. 2006b. Zmiany w intensywności eksploatacji i polityce zarybieniowej jeziorowych

Department of Fish Biology and Breeding, University of Warmia

Baikal, Kozgol or Imandra

Environmental preferences, current status and threats

Species protection and prospects

Department of Spatial and Environmental Economics / Department

In an appraisal of things, events or actions, certain values are assigned according to

Basic functions of lakes

The value of a lake as a resource of water for human uses

Lakes as real estate properties

Stagnant waters are

are marked with the sym-

The market value of lakes