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Author(s): Topi K. Lehtonen, David Gilljam, Lari Veneranta, Tapio Keskinen & Mikaela 
Bergenius Nord 
Title: The ecology and fishery of the vendace (Coregonus albula) in the Baltic Sea 
Year: 2023 
Version: Published version 
Copyright:   The Author(s) 2023 
Rights: CC BY 4.0 
Rights url: http://creativecommons.org/licenses/by/4.0/ 
 
Please cite the original version: 
Lehtonen, T. K., Gilljam, D., Veneranta, L., Keskinen, T., & Bergenius Nord, M. (2023). The ecology 
and fishery of the vendace (Coregonus albula) in the Baltic Sea. Journal of Fish Biology, 103(6), 
1463–1475. https://doi.org/10.1111/jfb.15542 
R E V I EW A R T I C L E
The ecology and fishery of the vendace (Coregonus albula) in
the Baltic Sea
Topi K. Lehtonen1 | David Gilljam2 | Lari Veneranta3 | Tapio Keskinen4 |
Mikaela Bergenius Nord5
1Natural Resources Institute Finland, Oulu,
Finland
2Swedish University of Agricultural Sciences,
Department of Aquatic Resources, Institute of
Coastal Research, Öregrund, Sweden
3Natural Resources Institute Finland, Vaasa,
Finland
4Natural Resources Institute Finland,
Jyväskylä, Finland
5Swedish University of Agricultural Sciences,
Department of Aquatic Resources, Institute of
Marine Research, Lysekil, Sweden
Correspondence
Topi K. Lehtonen, Natural Resources Institute
Finland, Oulu, Finland.
Email: topi.lehtonen@luke.fi
Abstract
Brackish water ecosystems often have high primary production, intermediate salin-
ities, and fluctuating physical conditions and therefore provide challenging environ-
ments for many of their inhabitants. This is especially true of the Baltic Sea, which is
a large body of brackish water under strong anthropogenic influence. One freshwater
species that is able to cope under these conditions in the northern Baltic Sea is the
vendace (Coregonus albula), a small salmonid fish. Here, we review the current knowl-
edge of its ecology and fishery in this brackish water environment. The literature
shows that, by competing for resources with other planktivores and being an impor-
tant prey for a range of larger species, C. albula plays a notable role in the northern
Baltic Sea ecosystem. It also sustains significant fisheries in the coastal waters of
Sweden and Finland. We identify the need to better understand these C. albula popu-
lations in terms of the predator–prey interactions, distributions of anadromous and
sea spawning populations and other putative (eco)morphs, extent of gene exchange
between the populations, and effects of climate change on their future. In this regard,
we recommend strengthening C. albula-related research and management efforts by
improved collaboration and coordination between research institutions, other gov-
ernmental agencies, and fishers, as well as by harmonization of fishery policies across
national borders.
K E YWORD S
environmental change, fishing, phenotypic plasticity, population, salinity, Salmonidae
1 | INTRODUCTION TO BRACKISH WATER
ENVIRONMENTS
This review covers the current knowledge of the ecology and fishery
of a freshwater salmonid fish, the vendace (Coregonus albula L. 1758),
in the brackish waters of the northern Baltic Sea. We place this topic
within a more general framework of fish communities and fisheries in
brackish waters, and the challenges that such environments induce to
freshwater fish in general and C. albula in particular. The aim is also
to elucidate the current state of these C. albula populations and their
fisheries, and to form predictions about their future by covering rele-
vant literature and official fishery statistics. This information is useful,
for example, when deciding on how to adjust management measures
in response to the expected environmental changes. For the readers'
convenience, we also provide a glossary (Table 1) with explanations of
the key terms and geographic areas of this review.
Received: 6 March 2023 Revised: 25 August 2023 Accepted: 26 August 2023
DOI: 10.1111/jfb.15542
FISH
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2023 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.
J Fish Biol. 2023;103:1463–1475. wileyonlinelibrary.com/journal/jfb 1463
Brackish waters are often defined by their medium levels of salinity
(Table 1). These aquatic environments can be found around the globe,
typically where waters from sources of low and high salinity meet
(Elliott & McLusky, 2002; Pérez-Ruzafa, Marcos, Pérez-Ruzafa & Pérez-
Marcos, 2011). Such conditions in, for example, river deltas, lagoons, and
estuaries, frequently promote high primary productivity (Correll, 1978;
Houde & Rutherford, 1993; Pérez-Ruzafa, Marcos & Pérez-Ruzafa, 2011).
Indeed, many brackish waters function as nursery areas or otherwise sus-
tain large biomasses of fishes that tolerate their varying environmental
conditions (Houde & Rutherford, 1993; Whitfield, 2016), commonly sup-
porting fisheries (Table 1) of high socioeconomic importance (Costanza
et al., 1997; Joyeux & Ward, 1998; Lamberth & Turpie, 2003; Pérez-
Ruzafa & Marcos, 2012). Nevertheless, only a few species live solely in
brackish water environments. Instead, these waters are commonly inhab-
ited by a mix of freshwater, marine, and anadromous (Table 1) taxa and
are typically dominated by a low number of species, often of marine ori-
gin (Beaudreau et al., 2022; Cabrera-Páez et al., 2021; Dyldin et al., 2020;
Thiel et al., 2003; Whitfield, 1999). The abundance and diversity of
marine species tend to increase, and those of freshwater species to
decrease, along gradients of increasing salinity (Guo et al., 2022; Kindong
et al., 2020; Morin et al., 1992; Thiel et al., 1995; Whitfield, 1999).
Because of the pronounced gradients of, and fluctuations in, physical
and chemical conditions of brackishwaters, they are considered to be nat-
urally highly stressed ecosystems (Elliott & Quintino, 2007; Teichert
et al., 2017). In addition, low average depths and close connectivity to the
adjacent terrestrial ecosystems havemademany brackishwaters and their
fish populations vulnerable to over-fishing (Haimovici & Cardoso, 2017;
Jackson et al., 2001; Ulman et al., 2020), climate change (Kashkooli
et al., 2017; MacKenzie et al., 2007), eutrophication (Karadurmus¸ &
Sari, 2022; Soria et al., 2022; Table 1), pollution (Barletta et al., 2019;
Islam & Tanaka, 2004), and species invasions (Daskalov &
Mamedov, 2007; Feyrer et al., 2003). These anthropogenic impacts have,
in recent decades, resulted in significant declines of economically and eco-
logically important brackish water fish populations in, for instance, the
Black Sea (Demirel et al., 2020; Oguz, 2017), Caspian Sea (Daskalov &
Mamedov, 2007), Marmara Sea (Demirel et al., 2022), brackish lakes and
lagoons (Haimovici & Cardoso, 2017; Mohanty et al., 2009), and certain
major river estuaries (Shan et al., 2013; Zhou et al., 2019). Hence, many of
these brackish water areas (and their fisheries) are in urgent need of effec-
tive recovery andmanagement plans and actions.
2 | THE BALTIC SEA AS AN ENVIRONMENT
FOR FRESHWATER FISH
The Baltic Sea (Figure 1; Table 1) is one of the world's largest brackish
water areas, and it shares many challenges with other major
brackish water bodies. Notably, it is among the most human-
TABLE 1 Glossary
Anadromous A fish migrating from the sea to a river to spawn
Baltic Sea Relatively shallow brackish water sea, a component of the Atlantic Ocean and enclosed by the land masses of Denmark,
Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, and Sweden
Bothnian Bay The northernmost part of the Gulf of Bothnia (see below), north of ~63
32'N (Finland) / ~63
59'N (Sweden), see Figure 1
Brackish water Often defined as water with salinity between 0.5 and 30 ppt
By‐catch Individuals of an aquatic species caught unintentionally while targeting other species or sizes of aquatic wildlife
Coregonid A fish of the subfamily Coregoninae in the family Salmonidae (see below)
Demersal Living (or taking place) near the bottom of a body of water
Discard The part of a catch that is not retained on board during commercial fishing
Eutrophication The process by which a water body becomes progressively enriched with nutrients, particularly nitrogen and phosphorus,
resulting in increased phytoplankton productivity
Fishery The enterprise of harvesting (or raising) fish and other aquatic life
Gulf of Bothnia The northernmost part of the Baltic Sea, between ~59
50'N and 65
54N, consisting of the Bothnian Bay and Bothnian Sea,
see Figure 1
Gulf of Finland The easternmost extension of the Baltic Sea, with Finland to the north, Estonia to the south and Russia to the east, see
Figure 1
Hypoxia The state of a low or depleted oxygen in a water body
ICES Statistical
Rectangles
A latitude‐longitude based area mapping system that covers the north‐east Atlantic, including the Baltic Sea, developed by the
International Council for the Exploration of the Sea (ICES)
PSU Practical Salinity Unit: a standardised way of measuring salinity of a water sample at 15 
C that, under most conditions, is
nearly identical with salinity measures 'ppt', '‰' and '0.1%'
Salmonid A fish of the family Salmonidae, including trout, chars, whitefishes, graylings, taimens and lenoks
TAC Total Allowable Catch: a control measure that limits the maximum overall quantity of the catch of one or multiple target
species during a set timeframe
Vendace species
complex
Coregonus albula and its closest relatives that some authors consider as conspecifics, including C. sardinella, C. vandesius, C.
trybomi, C. fontanae and C. lucinensis
Year‐class strength Usually defined as the number of fish spawned or hatched in a given year
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influenced and stressed seas in the world (Aps & Lassen, 2010;
Elmgren et al., 2015; Fleming-Lehtinen et al., 2015; Leppäkoski
et al., 2002; Möllmann et al., 2009; Viitasalo & Bonsdorff, 2022). It is
connected to the marine waters of the North Sea only through rela-
tively narrow and shallow passages at its south-western corner, mak-
ing it, in essence, a very large estuary, if not for its lack of significant
tides (McLusky, 1999; Pérez-Ruzafa, Marcos, Pérez-Ruzafa & Pérez-
Marcos, 2011). The influx of marine water to the sea is variable and
the overall turnover of its water mass is slow, resulting in a pro-
nounced salinity gradient and challenges related to eutrophication and
hypoxic (Table 1) conditions (Fleming-Lehtinen et al., 2015;
Viitasalo & Bonsdorff, 2022; Winsor et al., 2001). The salinity of the
sea stays mostly below 10 PSU (Table 1) and as low as at 2–3 PSU in
the Bothnian Bay and eastern Gulf of Finland (Figure 1; Table 1).
The relatively low and challengingly variable salinity levels of the
Baltic Sea have resulted in low species richness, yet a fish fauna that
consists of a unique mix of freshwater and marine species (MacKenzie
et al., 2007; Ojaveer & Kalejs, 2005; Olsson, 2019). The salinity varies
both among and within coastal locations, affecting fish distributions so
extensively that general models linking fish species diversity and salinity
have been based on the Baltic Sea data (Whitfield et al., 2012). Indeed,
many of the Baltic Sea fish populations live at the physiological limit of
their range (MacKenzie et al., 2007) and experience multiyear fluctua-
tions in abundance, which are, at least partly, triggered by changes in
environmental conditions or competitive interactions (Casini
et al., 2009; Lehtonen et al., 1993; MacKenzie et al., 2007; Ojaveer
et al., 2010). Interestingly, despite the young geological age of the Baltic
Sea (Björck, 1995), some of its inhabitants show signs of genetic adap-
tation to its environmental conditions or divergence between popula-
tions occupying different parts of the sea (Hill et al., 2019;
Johannesson & André, 2006; Leder et al., 2021; Wennerström et al.,
2017). For instance, in the southern Baltic Sea, European flounder (Pla-
tichthys flesus L. 1758) spawn pelagially (Table 1), whereas most of
those in the north spawn demersally (Table 1), with the two forms
showing a strong enough genetic divergence and reproductive isolation
(Momigliano et al., 2017) that the latter was recently described as a sep-
arate species, Platichthys solemdali (Momigliano, Denys, Jokinen & Mer-
ilä 2018).
Besides their intriguing ecological features, Baltic Sea fish populations
support several viable fisheries (Aps & Lassen, 2010; MacKenzie et al.,
2007; Zeller et al., 2011). The subfamily Coregoninae and other salmonids
(Table 1) are one such fish group, which has sustained recreational and
commercial fisheries of high socioeconomical importance, while being
particularly vulnerable to anthropogenic change (Dahlke et al., 2020;
Smialek et al., 2021). The Baltic Sea has both anadromous and resident
(i.e., sea spawning) salmonids (in genera Coregonus, Salmo, and Thymallus),
most of which have been important targets of fishing for hundreds of
years, if not for millennia (Lajus et al., 2013). While these fish have been
found to develop local differences in salinity tolerance (Fraser et al., 2011;
Larsen et al., 2008), the individuals that live outside their optimal salinity
(Arnesen et al., 1993) or temperature (Griffiths et al., 1992) ranges may
grow slowly and experience increased mortality.
F IGURE 1 (a) October 2021 mean salinity (PSU) in the Baltic Sea. Here, salinity is bounded to ≤10 PSU to clarify its gradient. Blue contours
highlight salinities of 2, 4, 6, and 8 PSU. (b) October 2021 mean temperature (degrees Celsius) at 1.5 m depth. The contours highlight 8, 10,
12, and 14
C. The different parts of the Baltic Sea, the Baltic Sea Proper in the south, the Gulf of Finland in the east, and the Bothnian Sea and
Bothnian Bay in the north (the latter two being collectively called the Gulf of Bothnia) are also shown. The salinity and temperature data originate
from the Swedish Meteorological and Hydrological Institute and have been modeled for the entire Baltic Sea by the ice-ocean model NEMO-
Nordic (https://doi.org/10.48670/moi-00013).
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3 | THE VENDACE IN THE BALTIC SEA
Coregonus albula is a small-sized (total length rarely exceeding 20 cm),
schooling planktivore that prefers cold water and matures early (Bøhn
et al., 2004; Gregersen et al., 2011; Lehtonen, 1981). Although both
juveniles and adults dominantly forage on zooplankton (Northcote &
Hammar, 2006; Sandlund, 1992; Viljanen, 1983), their diet is flexible,
with adults opportunistically predating on benthic crustaceans, insect
larvae, mollusks, and small fish, including conspecifics (Strelnikova &
Berezina, 2021; Urpanen et al., 2012). Their typical, but not exclusive,
living environments are large oligotrophic lakes in northern Europe,
including parts of Denmark, Estonia, Finland, Germany, Norway, Poland,
Russia, and Sweden. Both anadromous (Bogdanov et al., 2021) and sea
spawning (Björkvik et al., 2021; Enderlein, 1989; Veneranta et al., 2013)
populations reside the low-salinity waters of the northern Baltic Sea
(Gulf of Finland and Gulf of Bothnia; Figures 1 and 2). Locally, the spe-
cies may also be found, or was previously found, in more southern parts
of the sea (Lehtonen, 1981; Smitt, 1895). Similarly in lakes and the Bal-
tic Sea, C. albula spawn along the shores when the water has cooled in
October and November. The eggs then hatch the following spring
around the time the ice cover melts (Karjalainen et al., 2016; Koho
et al., 1991; Nyberg et al., 2001; Urpanen et al., 2005). The spawning is
F IGURE 2 Coregonus albula catches in different parts of the Swedish and Finnish waters of the Baltic Sea since 1998. (a) 1999–2001 (white
dots indicate additional Finnish coastal water areas where C. albula were commercially caught in 1980–1997), (b) 2002–2005, (c) 2006–2009,
(d) 2010–2013, (e) 2014–2017, and (f) 2018–2021. Colors indicate the yearly average commercial C. albula catch within each 50  50 km ICES
Statistical Rectangle (see Table 1) during the period stated in the panel. Note that in the Swedish fishery, effort and catch regulations (i.e., TAC,
see Table 1) affect the catches and therefore they do not necessarily correlate well with the abundance of the species.
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not concentrated within particular sites but takes place widely across
suitable areas (Björkvik et al., 2021; Karjalainen et al., 2018; Veneranta
et al., 2013). In the Bothnian Bay, important sites for reproduction are
located in both Swedish and Finnish coastal areas and river mouths,
where the fry are likely to remain for the first weeks of their lives after
hatching (Veneranta et al., 2013). Tagging experiments conducted in the
area suggest that C. albula return each year to the same region to
spawn, and then during summertime spread around wider areas to feed
(Enderlein, 1989). The typical maximum range of these migrations, how-
ever, seems to be only tens of kilometers (Enderlein, 1989).
Pronounced variability in year-class strength is typical of C. albula
populations, both in lakes (Karjalainen et al., 2000; Marjomäki, Auvinen
et al., 2021) and the Baltic Sea (Bergenius et al., 2013; Lehtonen, 1981).
This variation has been observed both as cyclic fluctuations (Marjomäki,
Auvinen et al., 2021) and more irregular alternations between strong and
weak year-classes (Axenrot & Degerman, 2016; Sarvala et al., 2020). The
postulated drivers of these oscillations include density-dependent sur-
vival of the youngest cohort, competition for food between cohorts, as
well as other intraspecific interactions (Hamrin & Persson, 1986; Marjo-
mäki Valkeajärvi et al., 2021). However, evidence also suggests that fish-
ing mortality (Sarvala et al., 2020), environmental factors (Auvinen
et al., 2004; Marjomäki et al., 2004), and interspecific interactions (see
below for details) can markedly affect the variation in population abun-
dance. Significant intraspecific and external factors are not mutually
exclusive but can have simultaneous and interacting effects on the pro-
nounced variation in C. albula abundance (Axenrot & Degerman, 2016;
Bergenius et al., 2013; Helminen & Sarvala, 1994), while high plasticity in
growth and fecundity probably dampen these fluctuations in the longer
term (Karjalainen et al., 2016).
Being a freshwater species, the spatial range of C. albula in the
Baltic Sea is presumably restricted by salinity more than any other sin-
gle factor (Enderlein, 1989; Lehtonen, 1981). Laboratory experiments
have shown that larvae are sensitive to salinities exceeding 5 PSU
(Jäger et al., 1981). Although larger juveniles and adult fish survive in
higher salinities (Jäger et al., 1981), the egg development is likely to
require even lower salinities (Veneranta et al., 2013). The presence
of additional physiological and ecological factors, such as tempera-
ture variation (Bergenius et al., 2013; Nyberg et al., 2001), eutro-
phication (Veneranta et al., 2013), intense intra- and interspecific
competition (Enderlein, 1981; Hansson, 1984), and predation
pressure, may also reduce the species' spatial range in the
sea. Indeed, these factors, together with the physiological
challenges of adapting to salinity, are among the typical limitations
to freshwater fish diversity in brackish water environments
(Whitfield, 2015).
Despite having only been able to establish within a limited spatial
range in the Baltic Sea, C. albula exhibits a significant potential to adapt
to local conditions. Sympatric forms that occupy different niches with
respect to the timing of spawning (Delling & Palm, 2019; Schulz &
Freyhof, 2003; Sendek, 2021), spawning migrations (Bogdanov
et al., 2021), body size and growth (Reshetnikov et al., 2020;
Strelnikova & Berezina, 2021), and diet (Strelnikova & Berezina, 2021)
have been documented both in lakes and the Baltic Sea. For example,
Strelnikova and Berezina (2021) reported the existence of small and
large-sized C. albula forms in the Gulf of Finland, with the latter occupy-
ing areas of deeper water. Moreover, while C. albula caught in the Gulf
of Finland are likely to be predominantly anadromous (Bogdanov
et al., 2021), those spawning in coastal waters seem to be dominating in
the Gulf of Bothnia (Enderlein, 1989; L
opez et al., 2022; Veneranta
et al., 2013). However, significant genetic differences between fish from
different parts of the Bothnian Bay indicate the potential presence of
local anadromous C. albula, besides coastal spawners (L
opez et al., 2022).
The occurrence of newly hatched C. albula fry in the lower reaches of
the river Tornionjoki that runs into the northernmost part of the Both-
nian Bay (Natural Resources Institute Finland, unpublished data) could
also indicate the presence of anadromous spawners in that river.
While the coastal spawners on the Swedish and Finnish sides of
the bay may be demographically separated (L
opez et al., 2022), cur-
rent knowledge of population boundaries and spawning migrations is
very limited and, as such, insufficient for the needs of knowledge-
based management plans. Therefore, additional spatial and temporal
sampling coverage is needed to better understand the distributions of
separate C. albula (sub)populations and the extent of gene exchange
between them. Interestingly, another species in the C. albula species
complex (Mehner et al., 2021; Sendek et al., 2013; Sendek, 2021;
Table 1), the closely related least cisco (Coregonus sardinella Valenci-
ennes 1848), which inhabits many North American and Siberian fresh-
waters, can successfully occupy estuary waters of varying salinities up
to 32 PSU (Craig, 1984). The populations assigned to each of these
two Coregonus species are not monophyletic and are so much alike
that some researchers argue that they constitute just one species
(Borovikova et al., 2013; Borovikova & Artamonova, 2021), further
suggesting a high potential for significant population-specific local
adaptations within the C. albula species complex (potentially including
the Baltic Sea populations). After having been introduced, C. albula
have also been able to rapidly invade new northern European river
and lake systems (Amundsen et al., 1999; Bøhn & Amundsen, 2001;
Kahilainen et al., 2011).
Coregonus albula is an important node in the Baltic Sea food web
because of its interactions with other species (and their fisheries). It
competes with other planktivores (e.g., young whitefish, Coregonus
lavaretus L. 1758, Baltic herring, Clupea harengus L. 1758, and smelt,
Osmerus eperlanus L. 1758) for food, with such competitive interac-
tions having potential to significantly affect the condition, and even
survival, of both C. albula and its competitors, especially during
periods of low food availability (Bøhn et al., 2008; Bøhn &
Amundsen, 2001; Enderlein, 1981; Hamrin & Persson, 1986; Nyberg
et al., 2001). Occasionally planktivorous fish (Miller et al., 1988),
including C. albula (Strelnikova & Berezina, 2021), can also be signifi-
cant predators of fish fry, which may, at least locally, affect the levels
of recruitment. Moreover, C. albula is an important prey of commer-
cially important predatory fish, including larger salmonids
(Heikinheimo, 2001; Hyvärinen & Huusko, 2005), and therefore, when
high in abundance, it has a positive effect on the survival of these cul-
turally and economically important species in the northern Baltic Sea
(Kallio-Nyberg et al., 2006). In the Gulf of Bothnia, it is also prey to
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gray seals (Halichoerus grypus Fabricius 1791) (Suuronen &
Lehtonen, 2012) and ringed seals (Pusa hispida Schreber 1775)
(Kauhala et al., 2019; Suuronen & Lehtonen, 2012). Indeed, in the Bal-
tic Sea, predation on C. albula by ringed seals can exceed the fishery
catches, potentially having a significant impact on the C. albula popu-
lations (Gilljam et al., unpublished; Hansson et al., 2018). Predation by
the European perch (Perca fluviatilis L. 1758) is also intense enough to
impact fluctuations of C. albula populations at least in a lake environ-
ment (Valkeajärvi & Marjomäki, 2004) and given that this predator is
predicted to benefit from climate change (Jeppesen et al., 2012;
Kokkonen et al., 2019) and has been increasing in commercial catches
(Official Statistics of Finland, 2023), its impact on C. albula in the Baltic
Sea may increase in the future. In contrast, if future trends also
include a continued decrease in salinity, Clupea harengus is likely to be
negatively affected (Polte et al., 2021), which can release additional
ecological space for C. albula.
4 | VENDACE FISHERIES IN THE
BALTIC SEA
4.1 | History of vendace fisheries in the Baltic Sea
Although relatively old records of C. albula fishery in the Baltic Sea's
brackish waters exist, the records are patchier than, for example,
those of the congeneric C. lavaretus (Bogdanov et al., 2021). The rea-
sons for the patchiness relate to a lower market value and extensive
abundance variations of C. albula (Bogdanov et al., 2021). Records
nevertheless show that it was a significant target species in the east-
ern Gulf of Finland in the 19th century (Bogdanov et al., 2021; Lajus
et al., 2013). Later, by the 1930s, this C. albula fishery had been con-
siderably reduced, presumably due to a decreased abundance, as a
result of (natural) changes in environmental variables, especially tem-
perature and salinity (Lajus et al., 2013). Starting from the latter part
of the 1940s, the importance of the C. albula fishery in the eastern
Gulf of Finland once again grew, with the Russian catches in this area
being as high as 1000 tonnes by the late 1950s (Bogdanov
et al., 2021; Lajus et al., 2013). The catches were still at a relatively
high level in the early 1970s, but have since then much decreased,
probably reflecting another period of a lower abundance due to both
anthropogenic pressures and natural changes in local conditions
(Bogdanov et al., 2021; Lajus et al., 2015).
Coregonus albula has, for decades, also been one of the most
important target species of commercial fisheries in the Gulf of Bothnia
in Sweden (Axenrot, 2021;Bergenius et al., 2018; Björkvik
et al., 2020) and Finland (Lehtonen, 1981; Official Statistics of Finland,
2023). The utilization of these northern stocks increased rapidly with
the use of commercial trawls, starting at the beginning and end of the
1960s in Sweden (Enderlein, 1978) and Finland (Lehtonen, 1981,
1983), respectively. The catches first peaked in the early 1970s
(Bothnian Bay: >1500 tonnes per year; Figure 3) and then decreased,
especially in Finland (Hildén et al., 1984; Lehtonen & Jokikokko, 1995;
Figure 3). Anecdotal reports (Lehtonen, 1981) suggest that during this
peak period the species was more widely distributed and harvested in
Finnish coastal waters than it has been since then. In Sweden, the C.
albula catches have remained at relatively high, albeit variable, levels
during most of the past 50 years (Figure 3), sustaining an economically
important fishery for roe (Bergenius et al., 2018; Björkvik et al., 2020).
4.2 | The current status of the Baltic Sea vendace
fisheries
In recent years, C. albula catches in the Russian part of the Gulf of
Finland have stayed at relatively stable but low (10 tonnes) levels
(Bogdanov et al., 2021). On the Finnish side of the gulf, the commer-
cial fishing effort has in recent decades been between low and nonex-
istent (Figure 2). In the Gulf of Bothnia, in turn, the current C. albula
fisheries operate mostly north of the N 63
 latitude (Figures 1 and 2).
On the Swedish side of the Gulf of Bothnia, the catches reached
almost 1700 tonnes in 2014–2015, which is the highest since modern
fishery started, but have decreased to less than 1000 tonnes during
the last few years (Bergenius, 2021; Figure 3). Catches in Sweden
have been intense enough to have a measurable effect on C. albula
recruitment, yet the impact of fishing on the population has been
smaller than that of the winter water temperature and salinity
combined (Bergenius et al., 2013). On the Finnish side, the commercial
C. albula catches have recently been increasing, reaching the highest
level since the 1970s, >500 tonnes, in 2022 (Figure 3).
In recent decades, trawling has been by far the most important fish-
ing method used by commercial C. albula fisheries in the northern Baltic
Sea (Finland 75%, Sweden 95%), followed by fish traps (fyke nets)
and gillnets (Figure 3). Gillnets dominate recreational C. albula catches
(>95% in Finland, Official Statistics of Finland, 2023). In Sweden, C.
albula are caught mainly for their highly valued roe, and only a small
proportion of the catch is consumed as fish meat. In particular, after roe
extraction, the remaining fish carcasses are either burned or used as ani-
mal feed. In Finland, the catch is used in a large part for human
0
500
1000
1500
2000
1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015 2020
Year
V
en
da
ce
 c
at
ch
 (
10
00
 k
g)
Gillnet Finland
Fyke Finland
Trawl Finland
Total Finland
Rest of Sweden
Trawl Sweden
F IGURE 3 Coregonus albula catches from the Gulf of Bothnia.
Total Finland data (1965–1979) are based on Lehtonen and Jokikokko
(1995). The 1980–2022 Finnish data are according to the Official
Statistics of Finland (2023). The Swedish data are according to
Lövgren et al. (2022).
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consumption as fish meat and increasingly also for roe and animal feed.
The current Baltic Sea C. albula fishery in Sweden is restricted to
40 trawl fishing licenses (Figure 4) and a fishing period of 5 weeks prior
to the spawning peaking in October. In addition, the Swedish Agency
for Marine and Water Management sets an annual catch quota after
considering the biological advice provided by the Swedish University of
Agricultural Sciences (Bergenius, 2021). Swedish gear restrictions to the
Baltic Sea C. albula fishery include a trawl modification requirement,
which prevents the catching of too-small individuals, and the size of
trawling vessels is limited to a maximum of 14 m. The allowed fishing
area is also restricted. Parallel with these general regulations, self-
regulation, such as additional area and time restrictions, is also encour-
aged. In contrast, in the Finnish waters of the Baltic Sea, the C. albula
fishing effort is currently not regulated. The profitability of the Finnish
C. albula fishery has had an increasing trend (Official Statistics of Fin-
land, 2023), while the highest trawling effort to date was reached in
2022 (Figure 5). Overall, a relatively large number of small vessels par-
ticipate in the Finnish fishery (Figure 4). The recreational C. albula fish-
ery in the Baltic Sea is important only locally, being much smaller than
the one in freshwaters (in recent years in Finland <50 tonnes per year
versus 700–2500 tonnes, respectively; Official Statistics of Finland,
2023). Note that the official catch statistics, reported above and in
Figures 2–6, are based on obligatory monthly catch reports by commer-
cial fishers. The fishers are required to report, among other things, the
gear they used, date and hours of fishing, fishing area (ICES Statistical
Rectangle, see Table 1) and the catch per species in kilograms. Data on
recreational catches are gathered less systematically and their estimates
are therefore more tentative. In absolute terms, the official statistics
may underestimate actual catches or have other inaccuracies, if the
catches are not duly reported by all fishers. We nevertheless expect the
statistics to capture temporal and other relative changes reasonably
accurately.
Regarding the ecological effects of the current fishery, it is also
relevant to consider the by-catch (Table 1), especially that of commer-
cial trawling (Figure 6), which is a key concern in fisheries manage-
ment and policy (Davies et al., 2009; Kennelly & Broadhurst, 2021).
The numbers of anadromous or sea-spawning ecotypes of C. lavaretus
caught as a by-catch of C. albula trawling can, at least occasionally, be
significant, warranting further assessment and monitoring (Leskelä &
Lehtonen, 1992; Marjomäki et al., 2016). Young individuals of other
larger species may also get caught by C. albula trawls (Jurvelius
et al., 2000). For example, sea trout (Salmo trutta L. 1758) smolts,
especially those that are reared in hatcheries to boost the threatened
populations of the Baltic Sea catchment area, may be vulnerable to
fishing practices that target other coastal species, including the sprat
(Sprattus sprattus L. 1758) and C. harengus (Degerman et al., 2012;
Kallio-Nyberg et al., 2007). Nevertheless, trout smolt mortality due to
the current commercial C. albula fishery seems to be low (Statistics
Database Natural Resources Finland, 2023; Figure 6). By-catch and
discard (Table 1) issues aside, as a fishing method, bottom trawling
can be very destructive to benthic habitats (Hiddink et al., 2017;
Thrush & Dayton, 2002). Coregonus albula fisheries, however, typically
use trawls over rocky habitats within restricted areas.
4.3 | The future of the Baltic Sea vendace
In the future, increasing temperatures in the Baltic Sea region are
likely to be particularly challenging to cold-adapted species, such as
C. albula and other native salmonids (Elliott & Bell, 2011; Graham &
Harrod, 2009; Karjalainen et al., 2014; Kumar et al., 2013). It is possi-
ble, albeit not certain, that the salinity of the Baltic Sea surface waters
will continue its recent declining trend with the warming climate
(Lehmann et al., 2022), which would stress the ecosystem (Lehmann
et al., 2022), but potentially benefit some freshwater species,
200
300
400
500
2000 2005 2010 2015 2020
Year
E
ffo
rt
 (
ho
ur
s 
of
 tr
a
w
lin
g)
F IGURE 5 Yearly fishing effort of commercial Coregonus albula
trawlers in Finnish coastal waters in 1998–2022 (Statistics Database
Natural Resources Institute Finland, 2023). Note the scale of the
y-axis.
50
100
150
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015 2020
Year
N
um
be
r 
of
 v
es
se
ls
>500 kg >1000 kg All catches Finland Sweden
F IGURE 4 Approximate number of active vessels fishing for
Coregonus albula. The Swedish vessel numbers are based on the
licenses for a 5-week C. albula fishing season in Bothnian Bay. In
Finland, the number of vessels is not restricted. The three Finnish
datasets are for vessels with any reported commercial C. albula
catches (including also small side catches when mainly targeting other
species) any time of the year anywhere on the Finnish coast (squares),
vessels with annual C. albula catches exceeding 500 kg (circles), and
vessels with catches exceeding 1000 kg (triangles). The Finnish
vessels are relatively small, with 89% and 3% of those reporting C.
albula catches in 2021 being <10m and >14 m, respectively (Statistics
Database Natural Resources Institute Finland, 2023).
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including C. albula (Pekcan-Hekim et al., 2016). However, because the
whole food web would be affected (Lehmann et al., 2022; Pekcan-
Hekim et al., 2016) and the projections for the changes in salinity,
stratification, and oxygen levels remain uncertain (Lehmann
et al., 2022; Viitasalo & Bonsdorff, 2022), the consequences for future
aquatic communities and their fish populations cannot be predicted
with any high level of certainty (Viitasalo & Bonsdorff, 2022). Because
C. albula populations in the Gulf of Bothnia and Gulf of Finland live at
the margin of their distributions, the increasing environmental varia-
tion may well impact them more (either positively or negatively) than
many other fish populations (Bergenius et al., 2013; Pekcan-Hekim
et al., 2016). Given these uncertainties, and the demonstrated effects
of environmental factors and fishing effort on C. albula recruitment
(Bergenius et al., 2013; Huusko & Hyvärinen, 2005), future manage-
ment measures should be set with caution to ensure that C. albula
catches stay within sustainable levels. In this respect, fish populations
are oblivious to national boundaries, highlighting the value of collabo-
ration between neighboring countries in research and management
efforts. This is especially important given that the current knowledge
of the population structure of the Baltic Sea Cor. albula, including the
extent of gene exchange between the Finnish and the Swedish parts
of the Bay, is incomplete (L
opez et al., 2022).
5 | CONCLUSIONS AND
RECOMMENDATIONS
Our literature review shows that the fish fauna in the brackish waters
of the Baltic Sea are vulnerable to human impact, including eutrophi-
cation, increasing temperatures (which could be coupled with decreas-
ing salinity), overexploitation, and habitat degradation (Elmgren
et al., 2015; MacKenzie et al., 2007; Viitasalo & Bonsdorff, 2022).
Commercially important species in the Baltic Sea include both anadro-
mous and resident salmonids. One of these, C. albula, occupies the
least saline parts of the Baltic Sea. While the species' distribution is
limited by salinity in combination with other factors (Bergenius
et al., 2013; Jäger et al., 1981), it also exhibits remarkable local adap-
tations and even sympatric forms that occupy slightly different niches
(Reshetnikov et al., 2020; Strelnikova & Berezina, 2021). The vendace
fishery is economically important on the Swedish side of the Bothnian
Bay, recovering on the Finnish side (Figures 2 and 3), and much smal-
ler in volume and predominantly recreational in the Gulf of Finland.
Since projections of future changes to the Baltic Sea area are consid-
erably uncertain, research efforts are needed to ensure appropriately
adjusted fisheries management measures.
Another pertinent research need is to understand the extent to
which the C. albula populations in the Baltic Sea (Gulf of Bothnia and
Gulf of Finland) are able to exchange genes with the adjacent freshwa-
ter populations, especially given that, in most rivers, dams block access
to the sea. In the same vein, we endorse unraveling the extent to which
the different C. albula populations migrate to rivers to reproduce
(i.e., are anadromous) versus completing their entire life cycle in the sea.
The two life-history strategies can be expected to differ in the likeli-
hood of gene exchange with other populations in the Baltic Sea and the
adjacent freshwaters. More investigations are also needed on the
extent to which the C. albula fisheries of the different nations surround-
ing the Gulf of Bothnia and Gulf of Finland are targeting shared versus
separate breeding populations (see L
opez et al., 2022). Earlier work
assumed that the Swedish coast of the Bothnian Bay would be a signifi-
cant source for adult C. albula on the Finnish side (Hildén et al., 1984),
whereas a subsequent tagging assessment suggested more localized
breeding populations (Enderlein, 1989). Further research is still needed
to map the coastal spawning areas, as well as levels of philopatry and
gene exchange by distance. Indeed, the current knowledge of the
139
232
1095
97
208
203
46
140
7
41 41
0
20
40
60
2000 2005 2010 2015 2020
Finland
Whitefish
Ruffe
Perch
Smelt
Others
23
41
20
5
14
4
2 4
15 22
23
20
15
9
7
4
7
6
7
10 21
21
36
32 37
0.0
0.5
1.0
1.5
2.0
2.5
2000 2005 2010 2015 2020
Sweden
Whitefish
Ruffe
Perch
Others
Year
%
 o
f v
en
da
ce
 c
at
ch
(a)
(b)
F IGURE 6 By-catches of the
Coregonus albula fishery in (a) Finland
and (b) Sweden. The percentage of the
most important by-catch species
compared to the catch of the target
species C. albula. ‘Others’ refers to all
other species. The percentages for the
most important by-catch, the Baltic
herring (Clupea harengus), are given as
numbers above the bars. Note that for
some fishers, especially in Finland, the
herring catches were a desired by-
product when C. albula was reported as
the main target species. The Finnish by-
catch data are based on limited samples
of the C. albula fishery. The Swedish
data are based on the trawl fishery,
which constitutes over 95% of the total
C. albula catch. Note the very different
scales for Finland and Sweden.
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species' population genetic structure in the Baltic Sea is still at best pat-
chy, despite the importance of such knowledge for informed manage-
ment and sustainable harvesting actions (Allendorf et al., 2008; Laikre
et al., 2005; Palsbøll et al., 2007; Wennerström et al., 2013). The effects
of the Bothnian Bay's planned wind power plant structures on the
C. albula and its fishery represent a related, timely knowledge gap.
Research, management, and policy-making parties should also
take into account that C. albula does not face environmental
changes in isolation but instead in interaction with other species
and, in many cases, their fisheries. In this regard, we endorse fur-
ther research on by-catches of the Baltic Sea fisheries. The research
efforts could, for instance, focus on whether C. lavaretus by-catches
of the C. albula fisheries are substantial enough to negatively affect
the various C. lavaretus populations and what practices could be
adopted to further reduce by-catch levels and mitigate their effects.
Such assessments should also be a part of the qualification process
of sustainable fishery certifications (Agnew, 2019; Björkvik
et al., 2020; Pappila & Tynkkynen, 2022; Pierucci et al., 2022).
More generally, we need a better understanding of the relative
impacts of the changing climate, eutrophication, habitat degrada-
tion, and fishing mortality. To attain these research and manage-
ment goals, we encourage intensified collaboration and
coordination efforts between fishers, research institutions, fisheries
management, and other governmental agencies, as well as harmoni-
zation of fishery policies among the countries surrounding the
northern Baltic Sea.
AUTHOR CONTRIBUTIONS
Topi K. Lehtonen conceived the study and wrote the first draft of the
manuscript. David Gilljam, Lari Veneranta, and Topi K. Lehtonen
designed the visualizations. All authors provided edits and additions
to the manuscript. All authors reviewed and accepted its final version.
ACKNOWLEDGMENTS
We thank Pirkko Söderkultalahti for help with finding the correct
Finnish statistics, Maria-Elena Bernal for proofreading an earlier ver-
sion of the text, and Bo Delling and an anonymous reviewer for con-
structive comments and suggestions.
ORCID
Topi K. Lehtonen https://orcid.org/0000-0002-1372-9509
Lari Veneranta https://orcid.org/0000-0001-5074-0822
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How to cite this article: Lehtonen, T. K., Gilljam, D.,
Veneranta, L., Keskinen, T., & Bergenius Nord, M. (2023). The
ecology and fishery of the vendace (Coregonus albula) in the
Baltic Sea. Journal of Fish Biology, 103(6), 1463–1475. https://
doi.org/10.1111/jfb.15542
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