Download Esox lucius: Northern pike

Esox lucius: Northern pike
Shannon Hennessey
FISH 423 Aquatic Invasion Ecology
Fall, 2011
Diagnostic information
Scientific name
Order: Esociformes
Family: Esocidae
Genus: Esox
Species: E. lucius
Common names:
Northern
pike,
pike,
jackfish, grand brochet
Figure 1: Northern pike Esox lucius color image (top) and black
and white image with diagnostic information (bottom) (Montana
Basic identification key
State Government’s Field Guide).
The Northern pike Esox lucius can
by Inskip (1982), all of which have specific
be characterized by a long body with a ‘duck
habitat requirements. The embryo stage is when
bill’ snout, a large mouth with many sharp teeth,
the fish has not yet emerged from the egg.
and cheeks that are completely scaled. The
Immediately after hatching, these become fry
dorsal and anal fins are set posteriorly on the
until they assume adult proportions at a size of
body with a narrow caudal peduncle and a
approximately 6.5 cm (Franklin and Smith
mildly forked caudal fin. E. lucius have greenish
1960). Juveniles are defined as those fish from
dorsal and lateral surfaces with yellow bars or
6.5 cm in length to the onset of sexual maturity,
rows of spots running along the sides, and a
or the stage at which their gonads begin to
whitish ventral surface (Figure 1). Body lengths
develop. Once these fish have reached sexual
range from 46-51 cm on average, with an
maturity, they become adults until the end of
average weight of 1.40 kg (Morrow 1980).
their life cycle. These fish are typically solitary
However, in the United States these fish can
and highly aggressive (Craig 2008).
grow to sizes up to 1.5 m long with weights up
Spawning occurs during the spring flood soon
to 35 kg.
after ice-out, when water temperatures are 812°C and can last from a few days to more than
Life history and basic ecology
a month (Franklin and Smith 1963, Inskip 1982,
Casselman and Lewis 1996). E. lucius migrate
Life cycle
up tributaries and spawns in calm, shallow
Esox lucius has a relatively simple life
waters of wetlands or flooded marshes where
cycle, in which they have 4 life stages as defined
there are mats of submerged vegetation. This
vegetation is essential not only to the survival of
Maturation is highly dependent on growth rate,
the eggs, but will also become important nursery
with males typically maturing at 34-42 cm total
habitat once the juveniles have emerged
length and females at 40-48 cm (Frost and
(Casselman and Lewis 1996, Mingelbier et al.
Kipling 1967; Priegel and Khron 1975). In the
2008). Eggs are deposited where they then stick
US, both males and females typically mature at
to the vegetation mats, which suspend them off
about 2 years of age, although there is variation
of the potentially anoxic sediments until
due to differential growing conditions (Inskip
hatching
later
1982). E. lucius can live as long as 24 years, but
(Franklin and Smith 1963, Casselman and Lewis
longevity is also strongly correlated with growth
1996). Water level is a very important factor in
rate (Miller and Kennedy 1948). At the more
embryo survival, due to the fact that E. lucius
southern end of their distribution, these fish may
usually spawn in water as shallow as 0.1 – 0.2 m
only live up to 4 years. (Buss and Miller 1961,
deep (Inskip 1982). High water levels during
Schryer et al. 1971, Inskip 1982). Adults are
spawning can also be associated with larger
also strongly associated with vegetation and
juveniles as the high waters bring an influx of
shallow waters, showing a preference for
nutrients, increasing the productivity in areas
submergent vegetation as it facilitates optimum
where larval fish are growing (Johnson 1957,
foraging efficiency (Casselman and Lewis 1996,
Casselman and Lewis 1996).
Craig 2008).
approximately
two
weeks
Once the eggs hatch, young-of-the-year fish
remain among the submerged vegetation, which
Feeding habits
provides important nursery habitat. At hatching,
E. lucius is characterized as a keystone
these fish are 7-9 mm in length on average
piscivore that can shape the composition,
(Frost and Kipling 1967, Forney 1968, Inskip
abundance and distribution of fish communities
1982). During the first few days, fry remain
(Craig 2008). They are visual predators that are
attached to the vegetation until their yolk sac
primarily active during the day, feeding on a
was absorbed (Frost and Kipling 1967, Inskip
wide variety of prey from invertebrates to fishes
1982). The juveniles grow rapidly, increasing
(Polyak 1957, Carlander and Cleary 1949,
their size and activity, and the shelter provided
Casselman and Lewis 1996, Craig 2008). As
by the vegetative cover helps enhance survival
opportunistic feeders E. lucius will feed on prey
(Casselman and Lewis 1996). As they grow,
that is seasonally abundant, although the size of
juveniles move to more floating and submergent
their prey is limited by both their size and the
vegetation, rather than the emergent vegetation
size of their prey (Craig 2008). They can
preferred by fry, but remain in shallow waters
consume a wide range of prey sizes, but have
(Casselman and Lewis 1996, Craig 2008).
also been shown to be size-selective predators
(Nilsson and Bronmark 2000, Craig 2008).
spawning groups, usually consisting of one
Although they can be cannibalistic, this has not
female and either one or several males. 5-60
been shown to comprise a large portion of their
eggs are released at any one spot, where they are
diet (Frost 1954, Lawler 1965, Mann 1976).
then fertilized by males (Svardson 1949, Clark
Aquatic vegetation is particularly important for
1950). The gametes are broadcasted, with no
feeding as these fish utilize and ‘ambush’ style
parental care provided post-spawning (Eddy and
of predation (Inskip 1982, Casselman and Lewis
Underhill
1996). Intermediate vegetation densities have
spawning group can release gametes over a large
been
predation
area of spawning habitat (Koshinsky 1979).
efficiency (Craig and Black 1986), and calm,
Franklin and Smith (1963) found the number of
clear waters provide better visibility associated
eggs produced to be highly dependent on female
with more efficient prey capture (Craig 2008).
body size, described by the equation y = 4401.4x
Fry begin feeding once they reach a length of
– 66245, with y equal to the number of eggs
approximately 10-12 mm in length, about 10
produced, and x as the total length of the fish in
days after hatching (Franklin and Smith 1963).
inches. Females weighing 0.75 lbs may produce
They initially feed on zooplankton, but soon
from 9,000 to 10,000 eggs, while a female
begin to feed on aquatic insect larvae and fish,
weighing 10 lbs may produce 100,000 eggs
marking the start of piscivory very early in their
(Lagler 1956).
associated
with
optimum
1974,
Inskip
1982).
Any
one
life (Hunt and Carbine 1951, Frost 1954, Inskip
1982). Within 4-5 weeks after hatching, at a
Environmental optima and tolerances
length of approximately 50-60 mm, their diet is
Northern pike are a cool water species,
comprised mostly of fish, and fry as small as 21
but have a wide range of environmental
mm have been shown to be cannibalistic (Hunt
tolerances
and Carbine 1951). As they continue to grow,
dissolved oxygen concentration, and water
their prey is predominantly fish, although they
clarity, which, in part, can be attributed to their
have been known to prey on crayfish, waterfowl
success as an invasive species (Casselman 1978,
and even small mammals (Frost 1954, Lagler
Steinberg 1992, Casselman and Lewis 1996).
1956, Seaburg and Moyle 1964, Lawler 1965,
These optima and tolerances, however, vary
Mann 1976, Inskip 1982).
among life-cycle stages (Inskip 1982).
including
water
temperature,
E. lucius embryos are tolerant of temperatures
Reproductive strategies
up to 19°C (Siefert et al. 1973), with maximum
E. lucius is an iteroparous species,
hatching success occurring at temperatures from
undergoing multiple reproductive events, with a
9-15°C and severe mortality at temperatures
relatively high fecundity. These fish form
near 5°C (Hassler 1970). Dissolved oxygen
concentrations of 4.5 mg/L is optimal for
shown no apparent stress at temperatures as low
embryo development, while concentrations of
as 0.1°C prior to freezing in a shallow lake
3.2 mg/L or less result in high embryo mortality
(Casselman
1978).
(Siefert et al. 1973). Embryos are particularly
temperature
tolerances,
sensitive to high rates of siltation (≥1 mm/day)
temperature can be lethal (Ash et al. 1974,
(Hassler 1970), and exposure to hydrogen
Inskip 1982). A significant decrease in feeding
sulfide of concentrations greater than 0.014-
has been observed in adults exposed to dissolved
0.018 ppm for 96 hours decreases hatching
oxygen concentrations around 2-3 mg/L, and
success (Adelman and Smith 1970a).
feeding has been shown to stop altogether at
Fry
survival
and
growth
is
greatest
Despite
these
sudden
wide
drops
in
at
concentrations less than 2 mg/L (Adelman and
temperatures from 18.0-20.8°C, with poor
Smith 1970b, Casselman and Lewis 1996). E.
survival at temperatures lower than 5.8°C and
lucius can inhabit areas with pHs ranging from
higher than 25.6°C (Lillelund 1966, Hokanson et
6.1-8.6 (Margenau et al. 1998), and can tolerate
al. 1973). Hydrogen sulfide concentrations
some salinity, having been found frequently in
greater than 0.004-0.006 ppm for 96 hours
the Baltic Sea (Crossman 1979).
significantly decreased survival of yolk sac fro
(Adelman and Smith 1970a).
Biotic associations
Juvenile E. lucius reach maximum growth rates
Biotic associations of E. lucius vary
at temperatures ranging from 19-21°C, with
across the native and introduced range of the
significantly decreased survival at temperatures
species, although there are numerous pathogens
lower than 3°C and above 28°C (Casselman
and parasites that can be associated with this
1978). Growth rates were hindered at dissolved
species. One notable ectocommensal organism
oxygen concentrations below 7 ppm, and very
associated
low juvenile survival occurred at concentrations
Capriniana piscium, also known as Trichophyra
below 3 ppm (Adelman and Smith 1970b).
piscium. C. piscium attaches to the lamellar
Feeding ceased at less than 2 ppm and no fish at
epithelia of the gill surface of E. lucius using
dissolved oxygen concentrations of 1 ppm
microfobrils that connect to the host cell
survived more than 20 hours in a study by Petit
membrane (Lom 1971, Hofer et al. 2005; Figure
(1973).
2). It was previously thought that high densities
Adults
have
the
greatest
with
E.
lucius
is
the
ciliate
environmental
of these organisms could cause tissue damage in
tolerances of the life-cycle stages, with a
the host, however, there is no evidence of C.
maximum temperature tolerance of over 30°C
piscium feeding on the host cells (Lom and
(Ridenhour 1957). Optimum growth occurs at
Dykova 1992). Instead, this commensal uses its
approximately 20-21°C, but adult E. lucius have
tentacles to feed on ciliates that are in the water
populations in 41 states, including Washington,
Oregon and Idaho (Figures 3 and 4).
In the Pacific Northwest, E. lucius has an
established population in Coeur d'Alene Lake,
Idaho, and has been reported to have established
populations in the Spokane River in both Idaho
and Washington, as well as Lake Pend Oreille,
Idaho, and the Pend Oreille River in Washington
and Idaho. There have also been general reports
of the presence of established E. lucius
populations in the Columbia River in Oregon as
well as Washington (Fuller 2011) (Figure 4).
Figure 2: C. piscium in the interlamellar space of fish
gills. The tentacles protrude from the gill surface where
History of invasiveness
they can capture prey (Hofer et al. 2005).
Esox lucius has a long history of
that passes through the fish gill lamellae (Lom
introduction and subsequent invasions, both in
1971, Dovgal 2002, Hofer et al. 2005).
the United States and elsewhere. This species is
historically absent from Mediterranean France,
Current geographic distribution
Distribution in the United States
and the PNW
In the United States, the native
range of Esox lucius includes the
Great
Lakes
and
Mississippi
River basins as well as Alaska,
Pennsylvania,
Missouri
and
Nebraska (Page and Burr 1991),
and
the
River
(Holton
South
Drainage
and
Saskatchewan
in
Montana
Johnson
1996).
However, E. lucius has been
Figure 3: Native and invaded ranges of E. lucius in the United States (US
reported
Geological Survey 2011).
to
have
non-native
Figure 4: Native and invaded range of E. lucius in the Pacific Northwest with general ranges as well as point
occurrences of populations (Adapted from the US Geological Survey 2011).
into
the
Hudson
River
Basin,
where
it
central Italy, southern and western Greece, and
subsequently spread to the lower Hudson River
northern
widely
by 1976 (Mills et al. 1997). Widespread
translocated throughout Europe (Kottelat and
stocking of E. lucius in the Chesapeake Basin
Freyhof 2007), introduced to Spain, Portugal,
began in the 1960’s in Pennsylvania, followed
and Ireland (Welcomme 1988), among other
by
locations. E. lucius has also been introduced to
(Denoncourt et al. 1975).
Algeria,
Morocco,
lucius fingerlings were released in the District of
Tunisia, and Uganda (Welcomme 1988, Lever
Columbia, although the survival of these fishes
1996).
is unknown (Christmas et al. 2000). In
As an important game fish, the introduction
Colorado, E. lucius was introduced to the Elk-
history of E. lucius outside of its native range in
head Reservoir of the Yampa River drainage in
the United States dates back to the 1800’s. The
1977, where they subsequently invaded the
first recorded introduction of Esox lucius in the
Green River basin of Colorado and Utah in 1981
United States outside of its native range was in
(Wick et al. 1985), with similar introduction
the 1840’s when the United States Fish
histories for other states with established E.
Commission intentionally introduced the fish
lucius populations.
Scotland,
Ethiopia,
but
has
been
Madagascar,
stocking
in
Maryland
and
Virginia
In 1992, 4000 E.
E. lucius was first introduced to the Pacific
for those stocking the water body, but this
Northwest in the early 1950’s (Brown 1971) and
unregulated introduction of E. lucius can play a
now occurs throughout the Columbia River
large role in facilitating the spread of this
basin. It was illegally stocked into Coeur
species (McMahon and Bennett 1996).
d'Alene Lake, Idaho, in the early 1970’s (Rich
In the Pacific Northwest, one of the initial
1993), where it then spread into several other
introductions of E. lucius was via illegal
lowland lakes in northern Idaho (McMahon and
stocking with Coeur d’Alene Lake, Idaho (Rich
Bennett 1996). Although E. lucius is native to
1993), resulting in the subsequent spread of this
Montana in the Saskatchewan River drainage, it
species
has now spread throughout the state (DosSantos
populations in Washington are a result of illegal
1991), and has also spread to Idaho, Washington
stocking in the Flathead, Bitterroot and Clark
and Oregon.
Fork River systems in Montana. These fish then
across
northern
Idaho.
E.
lucius
migrated into Lake Pend Orielle, through Idaho
Invasion process
by way of the Pend Orielle River, into
Washington (Washington Department of Fish
Pathways, vectors and routes of introduction
and Wildlife). In the past few years, these fish
The primary introduction pathway of Esox
have spread from Boundary Reservoir into the
lucius is the intentional stocking of the fish for
Columbia River, and can now be found in
recreational purposes (Fuller 2011). As an
Oregon as well, most likely though downstream
important sport fish, E. lucius has been
migration in the Columbia River (McMahon and
intentionally introduced across the country
Bennett 1996).
initially by the United States Fish Commission,
and later by various state agencies, to stimulate
Factors influencing establishment and spread
fishing opportunities in various parts of the
The establishment and spread of a species
country (Jenkins and Burkhead 1993; Scott and
is determined by the environmental tolerances of
Crossman 1973). Intentional introductions still
that species, as well as certain life-history
occur due to public pressures for fishing
characteristics. E. lucius can withstand a wide
opportunities, despite the risks associated with
range of environmental conditions (Casselman
introductions (McMahon and Bennett 1996).
1978, Steinberg 1992, Casselman and Lewis
Illegal stocking, or bait-bucket transfer, also
1996), increasing the likelihood of establishment
frequently occurs, in which E. lucius are
and spread of this species due to the fact that
intentionally introduced to an ecosystem without
there is a large amount of habitat with tolerable
authorization. This is typically done to promote
environmental conditions available. E. lucius is
fishing opportunities in the recipient ecosystem
also a relatively fecund species (Casselman and
Lewis 1996), which may allow for rapid
lucius populations can not only severely
establishment of populations and subsequent
decrease prey fish abundances (He and Kitchell
spread to new areas.
1990), but also create undesirable fisheries due
The main factor that may restrict the ability of E.
to their small size. If E. lucius populations were
lucius to establish and spread, however, is the
left unchecked, they could cause a loss of
availability of shallow habitat with submerged
revenue from sport fisheries as well as have
vegetation (DosSantos 1991) vital in spawning
drastic impacts on native fish populations
as well as adult foraging, as well as the presence
(Washington Department of Fish and Wildlife).
of suitable fluctuation of water levels during the
The effects that E. lucius can have on native
spawning period (see life-cycle section) (Inskip
fishes
1982). The nature of these specialized habitat
especially when considering the native fishes of
requirements therefore serve to limit the
the Pacific Northwest. One of the main concerns
distribution and likelihood of spread (Jones
associated with the spread of E. lucius is the
1990). Because these fish are highly predatory,
impact this species can have on local salmon
an
the recipient
populations (McMahon and Bennett 1996). E.
community is also necessary to support E. lucius
lucius have been shown to have drastic negative
populations (McMahon and Bennett 1996),
impacts on salmon populations as they feed on
which may further constrain the amount of
salmon smolts migrating to sea (Larson 1985,
suitable habitat available for colonization.
Petrvozvanskiy et al. 1988) and these fish have
abundant
prey
base in
have
very
important
implications,
been associated with the extinction of local
Potential ecological and/or economic impacts
There are many potential ecological and
economic
impacts
associated
with
the
salmonid populations (Bystrom et al. 2007,
Spens 2007, Spens and Ball 2008), making their
invasion a serious threat.
introduction of E. lucius outside of its native
While the introduction of E. lucius threatens to
range. As a keystone piscivore, this fish can
cause serious declines in a popular Pacific
shape
and
Northwest sport fish, E. lucius in itself is also an
distribution of fish communities (Craig 2008),
important sport fish, and there are strong
which
economic
the
has
composition,
important
abundance
implications
when
benefits
associated
with
the
considering the native fish communities, as well
introduction of this species. Stocking of these
as sport fisheries. In places where E. lucius is
fish can generate popular sport fisheries, which
overpopulated, there is a resultant overfeeding
can create important revenue that enhances local
on their prey base, which impedes the growth of
economies (McMahon and Bennett 1996).
these fish. Stunted growth, however, does not
However, this economic benefit can to lead to an
result in lower reproduction, so overabundant E.
increase in the frequency of illegal stocking
(Vashro 1990, 1995), further contributing to the
common
continued spread of E. lucius and the harmful
suppression and removal, although they tend to
effects that can follow.
be costly and/or labor intensive, with tradeoffs
strategies
for
fish
population
associated with each strategy (CDF&G et al.
Management strategies and control methods
2006).
An
important
management
strategy
in
There are several different management
preventing the spread of E. lucius is regulation
strategies and control methods for Esox lucius
of illegal introductions. Many states have laws
populations, the most effective of which,
that prevent the transport of live fish from a
however, is prevention. Once established, these
waterbody (e.g., Idaho, Oregon, and western
fish
habitats,
Montana), but there is strong opposition from
biotic
the public and it is difficult to closely manage
communities (He and Kitchell 1990, Craig
the actions of individuals stocking fish illegally
2008), making a preemptive response the best
(McMahon and Bennett 1996). In Montana, the
approach. One of the main prevention strategies
legislature stiffened penalties associated with the
is education, allowing for an early-warning
illegal stocking of fish, such as fines, suspension
system for new introductions in which a quick
of fishing privileges, and liability for cost of
response can be enacted, as well as increasing
restoring a fishery, and they have even offered
awareness of the impacts of E. lucius, hopefully
rewards for reporting such activity (McMahon
deterring the intentional introduction of the
and Bennett 1996). Following the actions of the
species.
Montana legislature in creating public incentive
However, in the cases where introduction has
seems to be a good place to start in terms of
already occurred, primary strategies for the
finding better ways to raise awareness and
control of E. lucius include the suppression of E.
prevent the introduction of invasive species.
can
drastically
spread
to
affecting
neighboring
the
recipient
lucius populations, the prevention of spread to
other waterways, and removal of as many
Literature cited
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Executive Director for Natural Resources
Kalispel Tribe
Other key sources of information and bibliog
Kalispel
raphies
P.O.
Usk,
Tribal
Box
39
WA
99180
(509)
445-1147
California Department of Fish and Game
Phone:
http://www.dfg.ca.gov/lakedavis/biology.html
Fax: (509) 445-1705
FishBase
Headquarters
Current research and management efforts
http://www.fishbase.org/summary/EsoxCurrently, there are numerous ongoing
lucius.html
efforts to control the spread of Esox lucius, with
Species
a focus on eradicating populations that have
Database. Pam Fuller. 2011. Esox lucius.
been introduced illegally. In Montana, E. lucius
http://nas.er.usgs.gov/queries/FactSheet.aspx?Sp
removal programs have been implemented in the
eciesID=676 Revision Date: 6/21/2010
Milltown Dam Reservoir to help restore native
USGS
Nonindigenous
Aquatic
cutthroat and bull trout. The dam impedes
salmonid migration causing them to concentrate
the Pend Orielle River. Surveys are being
in the reservoirs, providing easy prey for the E.
conducted to determine the relative abundances
lucius that reside there (Dickson 2003). The
of E. lucius, the structure of the overall fish
Montana Department of Fish, Wildlife, and
community, and the timing of E. lucius
Parks have temporarily lowered the later levels
spawning activities. The WDFW is attempting to
of the reservoir each year to kill yearling E.
learn more about the impacts of E. lucius on
lucius that occupy the back channels, resulting
native salmonid populations, using events that
in reduced fish populations, although the process
occurred in south-central Alaskan watersheds as
is labor intensive. Montana has also recruited
a model to attempt to prevent similar damage to
angler organizations to help control E. lucius
Washington salmonid populations, and has
populations,
proposed to remove E. lucius’s classification as
as
well
as
prevent
illegal
introductions (McMahon and Bennett 1996).
a game fish, causing it to be listed solely as a
The California Department of Fish and Game
prohibited species.
(CDF&G) has been treating a lake with an E.
lucius population with rotenone, a common fish
toxicant, in an attempt to eradicate the fish. The
first time this was done, E. lucius were found
two years later, so another eradication attempt is
planning to be undertaken (CDF&G et al. 2006).
Experimental control measures such as the use
of net barriers, electrofishing, and encircling
nets are also being used to control E. lucius
populations, as well as the blockage of spawning
areas, reducing food availability, increasing
public
awareness
and
implementing
a
comprehensive fish monitoring program.
In Washington, management efforts are being
directed towards minimizing the impacts of E.
lucius on native fish species and reducing E.
lucius abundances in the Box Canyon Reservoir.
The Washington Department of Fish and
Wildlife (WDFW) is working with the Kalispel
Tribe Natural Resources Department (KNRD) to
devise an appropriate management strategy for