Ecological Modelling 385 (2018) 58–64

Contents lists available at ScienceDirect

Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel

Partial exclusion of spawning Cyprinus carpio to improve coastal marsh T

habitat may come at the cost of increased carp population growth
⁎
Amanda Caskenettea, , Eva C. Endersa, Doug Watkinsona, Dale Wrubleskib
a
Fisheries and Oceans Canada, Canada
b
Ducks Unlimited Canada, Canada

A R T I C LE I N FO A B S T R A C T

Keywords: Common Carp (Cyprinus carpio) feeding and spawning behaviours, when abundant, negatively impact shallow
Carp exclusion water habitats essential for marsh ecosystem functioning. A common coastal marsh restoration practice is to
Cyprinus carpio create barriers to carp movement into a marsh from connected lakes. These barriers often do not exclude all carp
Delta Marsh from a marsh, and the lower adult density and improved habitat may increase overall carp recruitment success.
Coastal marsh restoration
A population model was created for Delta Marsh on Lake Manitoba, Canada. Varying the proportion of the total
Matrix population modelling
Wetland conservation
carp population spawning in the marsh indicates that, with partial carp exclusion, the improvement in spawning
conditions increases carp population growth rate and decreases generation time. Increasing fishing mortality can
help mitigate this effect, indicating that a multi-tiered management approach is best. It is important that fish
exclusion programs be implemented with awareness of this unanticipated result.

1. Introduction At Delta Marsh, a coastal wetland on the south shore of Lake

Common Carp (Cyprinus carpio, hereafter referred to as ‘carp’) are
one of the most widely distributed freshwater fish species in the world
(Zambrano et al., 2006). Carp often exist in very high abundances
where introduced, with detrimental ecosystem impacts (Bajer et al.,
2009; Kaemingk et al., 2017). Through feeding and spawning beha-
viour, carp disturb submerged and emergent vegetation (Lougheed
et al., 1998; Zambrano and Hinojosa, 1999), this may lead to phyto-
plankton blooms (Matsuzaki et al., 2007) and a turbid state (Lougheed
et al., 1998).
In efforts to restore marsh habitat, managers have implemented a
variety of carp control methods including poisoning (Schrage and
Downing, 2004), fertility control (Thresher et al., 2014), and reduction
of water levels after spawning (Yamamoto et al., 2006). A customary
control method is the use of fish exclusion screens that prevent the
movement of carp into their preferred spawning grounds (Meredith
et al., 2006). These control operations range from labour intensive
screened fishways where fish are hand sorted (e.g., Cootes Paradise
Marsh, Canada (Lougheed and Chow-Fraser, 2001)) to unmanned per-
manent screens placed in culverts preventing movement of all fish with
maximum body widths greater than the screen bar spacings, including
native fish (e.g., Murray–Darling Basin, Australia (Hillyard et al.,
2010)).
Manitoba, Canada, screens that prevent the movement of large carp in
Lake Manitoba, where they overwinter, from entering into Delta Marsh
are deployed at the beginning of the carp spawning migration and are
removed after most carp have spawned (Wrubleski et al., 2016). At
70 mm bar spacing, the screens should prevent migration of 99% of
carp > 630 mm in total length (French et al., 1999); however, in-
dividuals smaller than 630 mm and early migrants may still access the
marsh.
Carp have a protracted spawning period and are extremely fecund
(Stewart and Watkinson, 2004). The number of eggs increases with
maternal size, with the average carp producing hundreds of thousands
of eggs (Swee and McCrimmon, 1966). Prior to the establishment of the
Delta Marsh Carp Exclusion Project in 2013, Delta Marsh had minimal
submerged vegetation (Wrubleski et al., 2016) and an abundant adult
carp population. Carp recruitment is regulated by adult carp density,
predation, and larval habitat (Bajer and Sorensen, 2010; Bajer et al.,
2012; Weber and Brown, 2013). Most fish species that are known to
regulate carp abundance through the predation of eggs or larvae do not
exist in Delta Marsh (e.g., centrarchidae (Bajer et al., 2012)) or exist in
low abundance (e.g., Yellow Perch (Perca flavescens)). This implies that
carp recruitment is most likely regulated by availability of larval ha-
bitat and adult carp density. With such high adult carp densities, and a
deteriorated marsh, it is likely that very little successful carp

⁎
Corresponding author at: 501 University Crescent, Winnipeg, Manitoba, R3T-2N6, Canada.
E-mail addresses: amanda.caskenette@dfo-mpo.gc.ca (A. Caskenette), eva.enders@dfo-mpo.gc.ca (E.C. Enders), Doug.Watkinson@dfo-mpo.gc.ca (D. Watkinson),
d_wrubleski@ducks.ca (D. Wrubleski).

https://doi.org/10.1016/j.ecolmodel.2018.07.005
Received 1 December 2017; Received in revised form 14 May 2018; Accepted 7 July 2018
Available online 24 July 2018
0304-3800/ Crown Copyright © 2018 Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/BY-NC-ND/4.0/).
A. Caskenette et al. Ecological Modelling 385 (2018) 58–64

recruitment occurred in the marsh in most years. Since the Carp Ex-
clusion Project started, Delta Marsh has seen an increase in submersed
vegetation and decreases in adult carp abundance.
Reductions in the numbers of large adult carp have been shown to
increase water clarity, reduce nutrient levels and blue-green algal
blooms, contribute to changes in the benthic and zooplankton com-
munities, and increase the abundance of submersed aquatic macro-
phytes (Bonneau and Scarnecchia, 2014; Kaemingk et al., 2017). These
habitat changes can in turn result in increased recruitment of carp. Carp
spawn in shallow waters with abundant aquatic vegetation
(McCrimmon, 1968), and an increase in submersed macrophytes would
increase potential spawning habitat. These plants also provide habitat
for the early larvae, which attach themselves to the surface of the plants
soon after hatch (McCrimmon, 1968). Increased macrophytes means a
change in the zooplankton community and potential increase in larger-
sized zooplankton (Lougheed and Chow-Fraser, 2001), which are an
important food resource for larval carp (Lechelt and Bajer, 2016).
Clearer water and reduced turbidity may also contribute to increased
feeding efficiency by larval carp (Lechelt and Bajer, 2016).
Hence, with the resulting increase in the quantity the spawning and
quality of larval habitat for carp in Delta Marsh, along with a decrease
in adult carp abundance in the marsh, the likelihood of survival for carp
eggs spawned in the marsh may have improved since the Carp
Exclusion Project began. Monitoring of the fish community indicates
decreases in the abundance of large carp in the marsh but there is now
evidence of change in the abundance of young-of-the-year (YOY) carp.
Historically YOY carp, while caught in the marsh, have not been the
most abundant small fish species. The relative frequency of occurrence
ranged from third most common to rare, depending on the study and
water level (as reviewed by Parks (2006)). In 2016, however, small fish
sampling indicated that YOY carp were the most abundant fish species
caught, comprising over 57.5% of the total catch (St George, 2017).
The objective of this article is to determine how partial exclusion of Fig. 1. a) Diagram and b) matrix representing the elements of the stage-struc-
carp from a coastal marsh during spawning will affect carp population ture matrix population model for Common Carp (Cyprinus carpio) in Delta
level metrics if decreasing the numbers of adult carp in the marsh im- Marsh and Lake Manitoba. The model consists of four stages: a young-of-the-
proves carp spawning conditions. To achieve this goal, a matrix popu- year (YOY) stage, juvenile (J) stage, an adult stage that migrates into the marsh
to spawn (AMARSH), and an adult stage that remains in the lake year round
lation model was created and the resulting population level metrics
(ALAKE). Transition elements include the fecundity coefficient (F), the stage-
were compared across a range of carp exclusion efficiencies (hereafter
specific annual probability of remaining in stage i and probability of adults
referred to as ‘screen efficiency’), carp harvest rates, and the suitability returning to the marsh or the lake (Pi), and the transition probability of sur-
of spawning habitat outside of the protected marsh. We show that with viving stage i and moving to the next and probability of adults moving between
improved spawning conditions in the marsh, partial exclusion of carp the marsh and the lake (Gi).
will increase population growth rate and decrease generation time (i.e.,
how long it takes a population to grow by the anticipated quantity of
offspring produced over an individual’s lifetime (Caswell, 2001)). fi = H ηi φ 1 T (2)

2. Materials and methods
2.1. Model
The carp life cycle at Delta Marsh and Lake Manitoba is represented
by a post-breeding stage-structured projection matrix (Caswell, 2001).
The stage-structured matrix includes the fecundity coefficient (Fi), the
stage-specific annual probability of remaining in stage i (Pi) and the
transition probability of surviving stage i and moving to the next (Gi)
(Fig. 1).
Fecundity coefficients (Fi) represent the contribution of an adult to
next year’s YOY individuals. Since a post-breeding model is assumed,
the coefficient Fi includes the annual survival probability (σi), the stage-
specific annual number of female offspring per individual (fi), and the
probability of remaining in the stage (Pi) such that:
Fi = σifi Pi (1)
where fi is the product of the effect of the exclusion screens on survival
of eggs to the YOY stage (H), the average fertility (egg count) for a
female of stage i (ηi), the proportion of females in the population (φ),
and the inverse of the average spawning periodicity per year (T):
The probability of moving from i to i+1 (Gj) is defined as σi(γi) and
the probability of surviving and remaining in stage i (Pi) is defined as
σi(1-γi). For the YOY, γi was set to one as no individuals remained in
that stage for more than one year. For the juvenile and adult stages,
assuming that the age distribution within stages is stable (Lefkovitch,
1965), the term γi can be calculated as:
D D−1
γ =
( ) −( )
σi
λ
σi
λ
j D
( ) −1 σi
λ (3)
where λ is the largest eigenvalue of the matrix (Caswell, 2001) and is
set to 1 to represent a population at equilibrium, and D is the duration
of the stage (years). For juveniles, D represents the duration between
YOY and age at maturation. For adults, D represents the duration be-
tween age at maturation and the maximum age of the population (tmax).
In addition, the relative proportion of juveniles split between adult
stages (marsh or lake) and the relative proportion of the adults in the
marsh or in the lake depends on the efficiency of the screens (SE) to
prevent adult carp from entering the marsh.

59
A. Caskenette et al. Ecological Modelling 385 (2018) 58–64

Fig. 2. Map of Delta Marsh (highlighted on the insert with a white box), Canada; an 18,500 ha coastal wetland on the south shore of Lake Manitoba (50° 11′ N, 98°
19′ W) where screens (triangles) at all connections between the lake and the marsh control movement of Common Carp (Cyprinus carpio) between the marsh and the
lake.

2.2. Study system

Delta Marsh is an 18,500 ha freshwater coastal wetland located on

the south shore of Lake Manitoba (50° 11′ N, 98° 19′ W) in south-central
Manitoba, Canada (Fig. 2). It is a relatively shallow marsh, inhabited by
a diverse fish community (31 species) during the open water months
(Wrubleski et al., 2016). Exclusion screens are deployed in the spring
after the ice has melted in the channels and prevent most large fish from
entering the marsh to spawn. The exclusion screens are then removed in
August to allow large fish to move back and forth between the marsh
and the lake.

2.3. Parameter estimates

2.3.1. Individual growth

Estimates of growth were based on length-at-age data for carp col-
lected from Delta Marsh in 2009 (Weber et al. (2015), Fig. 3a). The von
Bertalanffy growth curve relates length and age:

FLt = L∞ (1−e−K (t − t0) ) (4)

where FLt is fork length at age t, L∞ is the asymptotic size, K is a growth

parameter, and t0 is the hypothetical age at which the fish would have a
length of 0 mm. In correspondence with Weber et al. (2015), t0 was set
to 0; the oldest recorded age of carp in Delta Marsh (tmax) was 21 years
(Weber et al., 2015).

2.3.2. Fecundity
In the absence of fecundity data for carp in Delta Marsh, values for
parameters ϑ and β (10−9.615 and 5.325, respectively) for the re-
lationship between the number of eggs (E) in relation to FL (mm):
E = ϑ FLβ
Fig. 3. a) Length-at-age data (points) for Common Carp (Cyprinus carpio) caught
in Delta Marsh, Canada in 2009 (Weber et al., 2015). Also included is the best
fit of the growth curve (solid black line) and 95% confidence intervals (dashed
lines). b) The estimate of Common Carp survival for each age (black line) and
(5) the point estimates (points) used in the model.

were adopted from Weber and Brown (2012) based on the lengths of 40
in the model was set to 5 years to be conservative.
carp from two lakes in eastern South Dakota, USA. Carp spawn yearly;
The effect of the exclusions screens on the survival of eggs to the
consequently, the spawning periodicity (T) was set to one. The popu-
YOY stage (H) is unknown for Delta Marsh. Badiou (2005) estimated
lation was assumed to be 36% female (Swee and McCrimmon, 1966).
that for Delta Marsh, every additional 100 kg/ha of carp in the marsh
The age at maturity for female carp in Delta Marsh is unknown, in si-
would result in a 2.9 g/m2 reduction in submerged vegetation biomass,
milar systems female carp mature anywhere from age 3 or 5 (Swee and
which relates directly to the amount of ideal larval carp habitat.
McCrimmon, 1966; Bajer and Sorensen, 2010); age at maturity for carp

60
A. Caskenette et al.
Fig. 4. Population growth rate (λ) for Common Carp (Cyprinus carpio) in Delta Marsh, Canada under a range of fishing mortalities (FM = 0.1, 0.3, 0.5, 0.7,
respectively), lake fecundity coefficients (HL), and screen efficiencies (SE) calculated using a simple stage-structure matrix population model. The contour lines
represent λ isoclines.
Similarly, Bajer et al. (2009) found that an increase in carp biomass was
strongly correlated with the decline in percent vegetative cover in two
shallow lakes in Illinois, USA. These studies suggest that higher screen
efficiency will result in an increase in carp spawning and larval habitat
in the marsh. In addition, with increased screen efficiency, the density
of carp in the marsh would decrease, alleviating density-dependent
effects on egg and larval survival. Accordingly, it was assumed that
realised fecundity scales directly to screen efficiency (SE) (as a proxy for
the number of carp in the marsh) with realised fecundity decreasing
proportionally with decreasing efficiency:
HM = c * SE
where HM is unit-less and represents the marsh fecundity coefficient,
i.e., the effect of screen efficiency on survival of eggs to the YOY stage
in the marsh and c is a scaling factor for HM. For the purpose of this
analysis, c was set to 0.1 to obtain a reasonable population growth rate
(λ) between 0.7 and 1.2. Changing the value for c does not affect the
qualitative interpretation of the results. For the purpose of this model,
we assume a linear relationship; however, the shape of the relationship
between SE and HM is unknown and should be determined empirically,
along with the value for c, if precise values for λ are desired.
While Delta Marsh is the largest spawning habitat for carp on Lake
Manitoba, there are tributaries and other coastal wetlands (Watchorn
et al., 2012) that provide spawning habitat. The effect of the spawning
habitat on survival of eggs to the YOY stage outside of the protected
marsh (lake fecundity coefficient, HL) was varied from 0.01 to 0.1 in the
model, paralleling HM, to determine the influence on the results.
2.3.3. Survival
Stage specific survival rates (σi) are unknown for carp. An equation
developed by Charnov et al. (2013) (Fig. 3b) was adopted to estimate
natural mortality (M) based on von Bertalanffy parameters FL, L∞, and
Ecological Modelling 385 (2018) 58–64
K:
−1.5
FL ⎞
M= ⎛ ⎜ ⎟ K
L
⎝ ∞⎠ (7)
This equation is a simplified version of an equation created by Gislason
et al. (2008) that was based on empirical estimates of natural mortality
for several species of fish, and has been used previously to represent
carp natural mortality (Lechelt and Bajer, 2016). For each stage,
average length (i.e., length at 0.5 years for YOY, and length of the
midpoint age for the juveniles and adults) was used for the respective
(6) mortality estimate (Fig. 3b). An accumulation of carp on the Lake
Manitoba side of the screens trying to enter into Delta Marsh provides
the opportunity for an effective commercial carp harvest. Four levels of
fishing mortality (FM) rates (i.e., low (FM = 0.1), medium (FM = 0.3),
high (FM = 0.5), and very high (FM = 0.7)) were compared to de-
termine the influence of carp harvest on the results. Total mortality (Zi
= M + FM) was then converted to survival (σ):
σi = e−Zi (8)
Since YOY survival could depend on availability of submerged ve-
getation (Lubinski et al., 1986), the YOY stage may be more appro-
priately represented with different survival rates for the marsh and for
Lake Manitoba, which may differ in extent, density, etc. of vegetative
cover. However, the relationship between YOY survival and submerged
vegetative cover is unknown, directionally consistent with the effect on
reproductive success, and so would most likely be captured in HM and
HL .
2.4. Model analysis
The population modelling and analyses were performed in R (R Core
61
A. Caskenette et al.
Team, 2015). Estimated population level parameters include carp po-
pulation growth rate (λ) and generation time (GT, years), that were
calculated from the population matrix with the package “popbio”
(Stubben and Milligan, 2007) according to Caswell (2001). For λ > 1
there is an increase in population growth, for λ = 1 the population is
stable, and for λ < 1 the population is declining in abundance.
3. Results
Carp population growth rate (λ), for low values of lake fecundity
coefficient (HL, ranging from 0 to 0.1) with increasing screen efficiency
(SE) will lead to an initial increase in λ of around 10% (Fig. 4). Growth
rate peaks at intermediate screen efficiencies, then declines to values
lower than that of no exclusion (SE = 0) at high screen efficincies. In-
creasing fishing intensities decreases the screen efficiency required to
decrease λ below that of no exclusion. Increasing fishing mortality also
decreases the overall maximum λ reached by the population. As HL
increases, the decline in λ is delayed to higher SE values. Model out-
comes suggest that the partial exclusion of carp from the marsh in-
creases population growth rate but can be mitigated somewhat by in-
creasing harvest.
Depending on SE, HL, and FM, simulated generation time for carp in
Delta Marsh ranges from 5 to 8.5 years. Generation time (GT) decreases
initially with increasing SE followed by an increase in GT at a SE around
90%, but only for the lowest HL values and fishing intensities (Fig. 5).
For medium and high HL, GT is consistently lower than what it would
be without the screens, with the lowest GT of 5 years at the highest FM.
The simulations suggest that partial exclusion of carp decreases the
generation time of carp.
Changing the model such that only the juveniles transitioning to the
adult stage can access the marsh, affected by screen efficiency (e.g., by
maturing at a size smaller than the screen excludes), produces similar
Fig. 5. Generation time (GT in years) for Common Carp (Cyprinus carpio) in Delta Marsh, Canada under a range of fishing mortalities (FM = 0.1, 0.3, 0.5, 0.7,
respectively), lake fecundity coefficients (HL), and screen efficiencies (SE) calculated using a simple stage-structured matrix population model. The contour lines
represent GT isoclines.
62
Ecological Modelling 385 (2018) 58–64
but less drastic changes in population growth rate and generation time
over increasing screen efficiencies (not shown). Increasing fishing in-
tensity has a larger mediating effect on population growth rate as all
adults are susceptible to harvest (not shown).
4. Discussion
Restoration practices that attempt to exclude spawning carp to
improve marsh habitats with partial carp exclusion may also improve
spawning and recruitment conditions for carp. The improved habitat
and lower densities of spawning carp in the marsh could affect carp
population dynamics. We demonstrated that partial exclusion can result
in a population growth rate that is higher than when there is no ex-
clusion, and is only lower than that of no exclusion when a substantial
proportion of the population is excluded. Identification of the qualita-
tive relationship between screen efficiency, population growth rate, and
generation time for carp will facilitate the development of informed
management strategies that may improve restoration of marsh habitats
and adjacent waterbodies.
In this study, we used a matrix population model, which was
parameterised to represent carp in Delta Marsh, Canada to determine
the effect of partial exclusion of carp from the marsh. Carp population
dynamics were simulated under the assumption that carp exclusion
resulted in improved recruitment success for carp in the marsh. An
increase in carp population growth rate and a decrease in generation
time were direct consequences of improved marsh habitat by partially
excluding carp from the marsh. To our knowledge, this is the first study
of its kind to examine how partial exclusion of carp from ideal
spawning habitats could affect carp spawning efficiency, in turn af-
fecting carp population dynamics. Brown and Gilligan (2014) similarly
analysed the effectiveness of control methods using a meta-population
model for carp. However, they did not consider the effect of improved
A. Caskenette et al.
spawning success on population growth rate from varying the propor-
tion of the population not susceptible to these control measures.
The interpretation of our model results are limited by assumptions
regarding foundational relationships in the matrix population model.
There are three main elements of a matrix population model: survival,
growth, and reproduction. In regards to survival, we assumed that
survival was related to the length of the individual, and that average
lengths for each stage were representative. While other methods of
calculating survival exist in the literature, we chose the length-based
method because it has previously been applied to carp (Lechelt and
Bajer, 2016) and is empirically supported (Gislason et al., 2008). In
addition, we also assumed for simplicity that the survival of the YOY
that spend their summer in the marsh would be the same as YOY that
are in the lake for the full year. YOY carp, however, prefer habitats with
dense submerged vegetation (Lubinski et al., 1986), which we assume
would increase in the marsh with higher screen efficiencies. The effect
of partial carp exclusion on YOY survival should, however, be direc-
tionally consistent with the effect on recruitment; with minimal effects
on the qualitative interpretation of the results.
Growth was represented in the matrix population model as the
transition between stages and did not vary across screen efficiencies in
the model. Factors that affect growth include population density and
habitat variables (Lorenzen and Enberg, 2002), which will vary with
screen efficiency. The most important transition, between the juvenile
and adult stages, depends on how juvenile carp are distributed between
the lake and marsh. The rate of carp return to Delta Marsh before they
are adults is unknown for Lake Manitoba, and so it would be difficult to
incorporate effects of partial exclusion on this stage transition, but
should be considered if applied to other systems.
Due to the lack of information, we assumed a linear relationship
between carp exclusion and the effect on reproduction in the marsh. It
is possible that this relationship would take other forms, and future
empirical studies should be performed to determine what they might
be. In addition, in our model recruitment is directly related to the
abundance of adult carp. Environment, however, is a strong predictor of
recruitment for some fish species (Lobón-Cerviá and Bradford, 2014;
Szuwalski et al., 2015), and may be important for carp (Phelps et al.,
2008; Weber and Brown, 2013). The inclusion of the marsh and lake
fecundity coefficients allows us to incorporate environmental effects on
recruitment; however, if this relationship with the environment is
strong enough to disentangle the stock-recruitment relationship, a more
complicated model may be required.
The linking of demographic parameters (e.g., growth rate, mortality
rate) to environmental variability (e.g., temperature, dissolved oxygen
concentration, water levels) increases fluctuations in population
abundance (Dixon et al., 1999). The effect of partial exclusion of carp
on population growth rate and generation time depended on the
spawning success outside of the protected marsh. At high screen effi-
ciency, when carp are not able to use Delta Marsh as a spawning
ground, reproduction from the carp population may depend on flooding
events that increase spawning habitat in the lake or other coastal
wetlands and tributaries around Lake Manitoba (King et al., 2003),
which, along with the predicted changes to population parameters, will
lead to increased population variability.
In conclusion, increased population growth and decreased genera-
tion time are interesting consequences of partial carp exclusion that
have not been considered before. Unintentional outcomes of increasing
or decreasing connectivity in aquatic systems can have profound effects
on the ecosystem; despite this they are often not considered and/or
incorporated in the management decision making process (McLaughlin
et al., 2013). It is important that wetland managers be aware of this
unanticipated result and should consider it when developing new fish
exclusion projects. Our experience at Delta Marsh demonstrates that
partial exclusion can be effective at improving marsh habitat. However,
a decrease in carp population size may not be achieved by partial ex-
clusion alone and multiple management strategies should be
63
Ecological Modelling 385 (2018) 58–64
implemented at once. What effect increased carp population growth
and decreased generation time will have on Lake Manitoba and its
coastal wetlands and tributaries is not known, but should be monitored
in the future.
Conflict of interest
None.
Acknowledgements
We thank Fisheries and Oceans Canada and the Institute of Wetland
and Waterfowl Research of Ducks Unlimited Canada for their support of
this work. Some of the data used was generated by the Restoring the
Tradition at Delta Marsh project. This project was funded by Ducks
Unlimited Canada, U.S. Fish and Wildlife Service through the North
American Wetlands Conservation Act, Manitoba Sustainable
Development, as well as grants and funds from numerous individuals
and foundations. We also would like to thank Bob Emery for his as-
sistance and proof reading, Susan Witherly for drafting the map, and
two anonymous reviewers for greatly improving this manuscript.
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