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The red king crab, Paralithodes camtschaticus, was intentionally transferred from Russian territorial waters in the Northern Pacific Ocean and introduced into the Barents Sea between 1961 and 1969 in order to create a new commercial fishery. A decade later a reproducing population was found to be well established in the latter region. The red king crab has since dispersed southwards along the coast of Northern Norway. Its ecological impacts on the native fauna have been investigated. From 2002 till 2007 the management of the commercial fishery has been undertaken jointly by Norway and Russia. Since then, management has continued within the countries respective fishery zones in the Barents Sea. In 2004 Norway was given free rein to apply all necessary management methods to limit the spread of the crab westwards of 26°E longitude.
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B.S. Galil et al. (eds.), In the Wrong Place - Alien Marine Crustaceans: Distribution,
Biology and Impacts, Invading Nature - Springer Series in Invasion Ecology 6,
DOI 10.1007/978-94-007-0591-3_18, © Springer Science+Business Media B.V. 2011
Abstract The red king crab, Paralithodes camtschaticus, was intentionally transferred
from Russian territorial waters in the Northern Pacific Ocean and introduced into
the Barents Sea between 1961 and 1969 in order to create a new commercial fish-
ery. A decade later a reproducing population was found to be well established in
the latter region. The red king crab has since dispersed southwards along the coast
of Northern Norway. Its ecological impacts on the native fauna have been inves-
tigated. From 2002 till 2007 the management of the commercial fishery has been
undertaken jointly by Norway and Russia. Since then, management has continued
within the countries respective fishery zones in the Barents Sea. In 2004 Norway
was given free rein to apply all necessary management methods to limit the spread
of the crab westwards of 26°E longitude.
1 Introduction
The red king crab Paralithodes camtschaticus (Tilesius, 1815) (Lithodidae
Samouelle, 1819) (Fig. 1) is among the world’s largest arthropods, reaching
~220 mm carapace length (CL), a weight over 10 kg (Powell and Nickerson 1965a,
Powell and Nickerson 1965b), and living up to 20 years (Kurata 1961).
It is native to the Northern Pacific Ocean (Fig. 2) with reported range from the
Korea and Japan, Kamchatka, the Aleutian Island chain, Alaska, and southeast to
Vancouver Island, Canada (Rodin 1990).
L.L. Jørgensen (*)
Institute of Marine Research, Tromsø, Norway
E.M. Nilssen
Department of Arctic and Marine Biology, Faculty of Biosciences, Fisheries and Economics,
University of Tromsø, N-9037 Tromsø, Norway
The Invasive History, Impact and Management
of the Red King Crab Paralithodes
camtschaticus off the Coast of Norway
Lis Lindal Jørgensen and Einar M. Nilssen
522 L.L. Jørgensen and E.M. Nilssen
Fig. 1 Dorsal view of Paralithodes camtschaticus (photographer: Lis Lindal Jørgensen, Institute
of Marine Research)
Fig. 2 The native distribution of the red king crab (yellow colour) along the coasts of Korea,
Japan, Russia, Alaska, and Canada
523The Invasive History, Impact and Management of the Red King Crab
The red king crab was collected by Russian scientists during the 1960s and
1970s from Peter the Great Bay, Okhotsk Sea, and introduced into the Barents Sea
(Orlov and Karpevich 1965; Orlov and Ivanov 1978) (Fig. 3). Between 1961 and
1969, 1.5 million first stage zoeae, 10,000 1–3 year old juveniles (50% females and
50% males) and 2,609 5–15 year old adult (1,655 females and 954 males) crabs
from West Kamchatka, were intentionally released into the Kolafjord, east Barents
Sea, Russia, in order to create a commercial fishery (Orlov and Karpevich 1965;
Orlov and Ivanov 1978). In the Russian part of the Barents Sea the highest densities
were observed on both sides of the Rybachi Island (Fig. 4) during late 1980s and
early 1990s. Later in the 1990s, the red king crabs became abundant along the
eastern part of the Kola Peninsula and were reported from Cape Kanin and the
entrance of the White Sea during 2002. Further northwards the crab was found on
the Kanin Bank and at the Goose Bank (Zelina et al. 2008).
Fig. 3 Red king crab dispersal in the Barents Sea. Embedded map showing the translocation of
crabs from West Kamchatka, North Pacific Ocean westwards into Kolafjord (see fig. 4), east
Barents Sea
524 L.L. Jørgensen and E.M. Nilssen
In 1992 the red king crab became abundant in Norwegian waters, initially
reported from southern Varangerfjord (Fig. 4). By 1994 P. camtschaticus spread to
the northern side of the fjord. The crab has increased fourfold in Varangerfjorden
within 12 years (Table 1). In 1995 it was recorded in Tanafjord and the population
has been relatively stable in the period 1999–2007 (Table 1). Further range exten-
sions were noted in Laksefjord and Porsangerfjord during 2000, and by 2001 several
adult crabs were caught west of Sørøya and west of the North Cape. In 2002 the crab
were captured close to Hammerfest and three specimens were recorded about 120
nautical miles west off the North Cape (Hjelset et al. 2003; Sundet 2008).
The crab population along the northern coast of Norway was estimated to num-
ber 2.9 million individuals in 2001 and 3.5 million in 2003 (Hjelset et al. 2003). In
2007 the population in Norwegian waters was estimated at 4–5 million individuals
(Sundet 2008). That number is an underestimate as only individuals with a carapace
longer than 70 mm and at water deeper than 100 m are included.
Fig. 4 The spreading of the red king crab along the northern coast of Norway
525The Invasive History, Impact and Management of the Red King Crab
2 Spreading, Settling, Podding and Migration
The larvae of the red king crab develop in the coastal zone. In the 2 months after
hatching, the pelagic larval stages can be transported by currents considerable dis-
tances (Pedersen et al. 2006). This period must be synchronised with the spring
phyto- and zooplankton peaks in the upper 15 m of the water column (Shirley and
Shirley 1989). The larvae settle in shallow waters (<20 m) on sponges, bryozoans and
macroalgae (Marukawa 1933). Successful recruitment depends on a well-developed
sessile community with extensive areas of dense concentrations of hydroids, bryozo-
ans, and sponges needed to support a massive settlement of larvae.
Red king crabs smaller than 20 mm carapax length (CL) lives a cryptic and soli-
tary life, sheltering beneath rocks and stones and in crevices. In the second year
podding behaviour (Fig. 5) appears (Dew 1990). Podding is when the crabs congre-
gate in large, tightly packed groups (Powell 1974). The smallest and largest crabs
found in any pod are 24 and 69 mm CL, respectively. Pods therefore form during
the latter part of the second year, exist throughout the third year, and continue a
short time into the fourth (Powell and Nickerson 1965a, Powell and Nickerson
1965b). When the density of the crab approaches 6,000 individuals, pod structures
transforms into elongate piles and dome shaped piles do not commonly occur until
the fourth year when crabs are 60–97 mm CL (Powel and Nickerson 1965a). The
pods are held during the daytime, but disperse into a nightly foraging aggregation.
This was explained by changes in water temperature, crab weight, and time of
Table 1 Average catch per unit of effort (CPUE) (number of
crabs per trawl hour) with 95% CI (confidence interval) of the red
king crab from the scientific cruises in the period 1995–2007
(From Hjelset et al. 2009)
1995 10.5 ± 3.6
1996 19.1 ± 7.0
1997 21.0 ± 7.7
1998 13.7 ± 2.9
1999 17.4 ± 4.5 18.3 ± 9.7
2000 25.0 ± 13.3 5.2 ± 2.7
2001 20.5 ± 10.0 6.0 ± 2.8
2002 15.6 ± 5.8 18.9 ± 9.7 2.5 ± 4.9
2003 19.7 ± 7.5 38.8 ± 18.9 37.9 ± 71.7
2004 30.4 ± 17.2 25.8 ± 8.2 25.4 ± 39.1
2005 33.3 ± 21.9 23.5 ± 9.6 13.0 ± 16.0
2006 41.5 ± 25.4 31.0 ± 14.2 25.0 ± 31.0
2007 45.8 ± 25.7 24.8 ± 9.2 25.9 ± 19.1
Not available
526 L.L. Jørgensen and E.M. Nilssen
sunset by Dew (1990). A trend of increased foraging time and movement to deeper,
cooler water was apparent after mid-April, as water temperatures reached 4°C and
began a sustained summer increase (Dew 1990).
Immature crabs (CL<120 mm), generally remain along the coast at 20–50 m
depth (Wallace et al. 1949), and are seldom associated with adults in deep water.
Adults occur on sand and mud bottoms (Vinogradov 1969; Fukuhara 1985) and
aggregate according to size, life history group or sex. The adult crab undergoes
two migrations, a mating-moulting migration and a feeding migration (Fig. 6). The
patterns of behaviour are similar off the coasts of Japan, Russia, and Alaska
(Marukawa 1933; Powell and Reynolds 1965; Vinogradov 1969). The shoreward
migration to shallow waters (10–30 m) takes place in late winter and early spring
when the crabs mate, breed (Marukawa 1933; Wallace et al. 1949; Powell and
Nickerson 1965a, b) and hatch their eggs (Stone et al. 1992). Extensive aggrega-
tions of both sexes occur during the spring spawning season. These spawning
aggregations may also be found also in shallow water where kelp occurs (Powell
and Nickerson 1965a, b). The kelp may provide shelter for the females following
moulting ecdysis, and during mating (Jewett and Onuf 1988). Spawning is fol-
lowed by migratory feeding movements, of both sexes, towards progressively
deeper water (300 m). After this period, the sexes form separate aggregations for
the remainder of the year (Fukuhara 1985), and are not found together until the
following mating season (Cunningham 1969).
In Russian waters the crab occurs both along the coast and offshore, while in
Norwegian waters, the crab is distributed solely along the coastline (Fig. 3). Since
along the Russian coast the bottom slopes gradually, whereas in the Norwegian
Fig. 5 Podding of juvenile red king crab (Paralithodes camtschaticus) in Norwegian fjord (Photographer:
Geir Randby, Lillehammer Film)
527The Invasive History, Impact and Management of the Red King Crab
fjords the bottom descends abruptly to deep water (300 m), it is proposed that the
pattern of distribution is dependent on the coastal topography. This gently sloping
coastal topography is also found in the north Pacific habitats, where the crab
migrates far from the coast to reach deep water. The steeper topography may keep
the Norwegian population close to the coast or inside the fjords year round.
3 Temperature Tolerance
The red king crab tolerates temperatures from −1.7 to at least +15°C (Rodin 1990),
these tolerance limits vary at different stages of its life history. Temperature prefer-
ences of immature crabs (50–100 mm CL) are at <3°C as determined in laboratory
studies (Hansen 2002). In the Barents Sea and the northern Norwegian Sea the
temperature at 100 m depth in winter varies from 0°C to ~+6°C. Recently, it has
been experimentally demonstrated that larval survival is affected by the water
temperature in which the egg carrying females had been kept (Sparboe pers. comm.).
Females acclimated to 14°C produced larvae with higher survival rates at high tem-
perature compared with larvae from females acclimated to 4°C and 8°C. Survival
was high (almost no mortality) for all crabs exposed to challenge temperatures
from −1.7°C to 15°C independent of acclimation temperatures (4°C , 8°C and 14°C)
Fig. 6 Seasonal migration of Paralithodes camtschaticus: the mating-moulting migration in the
spring/summer period to various substrates with benthic communities principally composed of
calcified prey organisms, and a subsequent feeding migration in winter/autumn to soft substrate
where annelids occur (inset: juvenile red king crabs associated with kelp)
528 L.L. Jørgensen and E.M. Nilssen
(Sparboe pers. comm.). This result may indicate that the red king crab may
successfully invade also more southern habitats along the Norwegian coast (Larsen
1996; Sparboe pers. comm.).
The population of West Kamchatka overwinters on the continental slope where
the warmer Pacific Ocean water mixes with the colder waters of the shallow shelf.
The migration from the over wintering area to shallow water depends on bottom
water temperatures, as well as the physiological conditioning prior to spawning and
moulting (Rodin 1990). Large numbers of adult crabs assemble in shallow waters
(10–15 m) in May–June when temperatures are approximately 2°C. Following
reproduction in June and July, adults forage at around 50 m depth where the water
is 2°C. Once temperatures decrease, the crabs disperse to deeper water for overwin-
tering (Rodin 1990).
Amazingly, a single red king crab male was recorded in the comparatively
“warm” Mediterranean Sea, though no explanation is given of its mode of introduc-
tion and survival so far south (Faccia et al. 2009).
4 Food and Feeding
The crab’s food preference varies with age and stage. The pelagic larvae feed on
both phytoplankton and zooplankton (Bright 1967). Once settled, the juveniles feed
on hydroids, the dominant component of the epifauna on the Kamchatka shelf
(Tsalkina 1969). Dew (1990) reported that young crabs (CL > 20 mm) feed on sea
stars, kelp, Ulva spp., red king crab exuviate, bivalves of the genera Protothaca and
Mytilus, nudibranch egg masses, and barnacles. Occasionally, crabs were observed
dragging around large sea stars during the nocturnal foraging period. These stars
were sometimes left near the base of the pod in the morning, and taken up again
upon pod break-up. Adults are opportunistic, omnivorous feeders (Cunningham
1969). They feed on the most abundant benthic organisms, though usually one food
group/species dominate their diet and this varies regionally (Kun and Mikulich
1954; Kulichkova 1955; Jewett et al. 1989). Most common food items are echinoderms
(Ophiura spp., Strongylocentrotus spp.) and molluscs (Nuculana spp., Clinocardium
spp., buccinid and trochid snails) (Cunningham 1969). Calcareous-shelled food
items are more frequent in the diet of post-moult crabs (Herrick 1909; Fenyuk
1945; Logvinovich 1945). Kulichkova (1955) suggested that crabs need to replace
calcium carbonate lost during moulting and that the young clams and barnacles in
shallow waters fulfill this need. At times of moulting, growth and reproduction, the
food intake declines but such pauses do not normally last more than 2–3 weeks
(Kulichkova 1955) and thereafter the crabs feed avidly (Takeuchi 1967). The crabs
feed on bivalves and echinoderms during spring and summer months when in shal-
low areas, and polychaetes in autumn and winter where they migrate to deeper
water (Gerasimova 1997). Crabs contain significantly more food in their guts dur-
ing spring-early summer (Takeuchi 1967; Jewett et al. 1989) when compared with
the late summer-autumn-winter (Jewett and Feder 1982).
529The Invasive History, Impact and Management of the Red King Crab
Adult crabs feed either by grasping and tearing apart larger invertebrates or by
scooping sediment by the lesser chela and sieving it through the third maxillipeds.
Scooping sand was often observed by Cunningham (1969) during periods when no
larger food was immediately available. Logvinovich (1945) referred to the frequent
presence of sediment in the stomachs and intestines of crabs. Foraminifera, minute
molluscs and amphipods found in stomach contents probably result from feeding
by sieving, as these either burrow in or occur on sediments. Logvinovich (1945)
suggested this as an alternative method of feeding when larger prey is unavailable.
Observations on the degree of gut fullness would indicate that crabs browse on food
as it is encountered (Cunningham 1969). Calculations indicate that a young adult
crab consumes 6 g, and juvenile crab 1.7 g within 25 h at 3°C, and 16 g and 3.5 g
respectively at 6°C (Jørgensen et al. 2004). Laboratory studies indicate a daily
ingestion rate of more than 70 g (squid) for young adult crabs at 5–9.4°C (Zhou
et al. 1998). Pavlova et al. (2007) showed that juveniles consume a mixture of
polychaetes, bivalves, ophiuroids, echinoids, asteroids weighing 0.7–26 g daily,
based on soft tissues. However, identification of prey items and calculation of their
weight from gut contents is inaccurate because decapods rarely swallow prey
whole, rather they tear it apart. These fragments are shredded further in the gastric
mill and are mostly unidentifiable. If to the weight of consumed soft tissue are
added the undigested shells (Chlamys islandica, Strongylocentrotus droebachien-
sis, Modiolus modiolus, Astarte sp., Buccinum undatum, Asterias sp. or Henricia
sp.) mature and immature crab show a daily foraging rate (killing or mortally dam-
aging) between 150 and 300 g at 5–6°C (Jørgensen 2005; Jørgensen and Primicerio
2007), 17–408 g when feeding solely on scallops within 24 h (Anisimova et al.
2005; Jørgensen and Primicerio 2007), and 1–101 g per 24 h when feeding on sea
urchins (Gudimov et al. 2003; Jørgensen and Primicerio 2007).
The above results might indicate a range from “low” (high abundance of prey,
high species richness, prey of low foraging preferences, or not foraged benthic
species) to a “strong” (low abundances of prey, species richness is low, highly pre-
ferred and flat-bodied prey species) impact on native local communities depending
on the abundance of prey and the number of red king crabs. Because food appears
to be the sole factor that could limit the increase in red king crabs numbers within
the Southern Barents Sea (Gerasimova 1997), it is most likely that the invasive spe-
cies, particularly in high abundances, will have a measurable effect on native prey
5 Ecological Impact
There is a growing recognition that aliens may interact negatively with the native
species in the recipient communities (e.g., Elton 1958; Lodge 1993; Carlton 1996;
Ruiz et al. 1997; Walton et al. 2002; Ross et al. 2003). Due to the body size, long
life span, predaceous behaviour, large population size and rapid dispersal of the red
king crab, questions have been raised as to its impact on the native benthic community.
530 L.L. Jørgensen and E.M. Nilssen
Since the establishment of the crab in the Barents Sea, studies on its predatory
effect have been undertaken (Sundet et al. 2000; Haugan 2004). The crab feeds on
a range of molluscs, sea urchins (Strongylocentrotus droebachiensis) and other
echinoderms, crabs, polychaetes, sipunculids and fish (Sundet et al. 2000). Indeed,
it was shown that some benthic taxa decreased considerably in abundance since its
introduction, and that changes have occurred in the benthic community structure in
the investigated fjords (Anisimova et al. 2005). It was calculated that the crab preys
upon 15% of the total coastal population of Strongylocentrotus urchins (Gudimov
et al. 2003; Pavlova 2009). Experiments of the potential impact of the invading crab
on the beds of the native scallop, Chlamys islandica, showed that the scallop had
no size refuge. The scallop’s flat shell is easily handled by both small and large
crabs (Jørgensen 2005; Jørgensen and Primicerio 2007), though small crabs seem
to prefer smaller scallops (Gudimov et al. 2003). Larger prey items with dome
shaped bodies, sponges, sea cucumbers and sea anemones were not preyed upon
(personal laboratory observations made by the author). Scallop beds with a rich
associated fauna are less vulnerable to predation than beds with few associated spe-
cies, had several possible prey items to forage in the rich species associated scallop
bed compared to the scallop bed with few other species than the scallop (Anisimova
et al. 2005; Jørgensen 2005; Jørgensen and Primicerio 2007).
Anisimova et al. (2005) calculated that the crab population consumes 37 tonnes of
capelin (Mallotus villosus Cuvier, 1829) eggs in a Barents Sea fjord during 3 months,
and extrapolated this value to the whole Barents Sea crab population. The study con-
cluded that the crab may impact 0.03% of the egg mass laid by the capelin.
In order to forecast possible impact in new or in already invaded areas, a study
of the quantitative values of the prey (killed or mortally damaged specimens) is
needed, and possible recipient areas need to be surveyed ahead of the crabs’ arrival.
The baseline surveys should include epifauna and infauna as the crab preys on
components of both.
6 Economic Impacts
The development of the crab fishery in Norway is illustrated in Table 2. The data
indicate that from 1994 to 2007 the total allowable catch (TAC) and effort increased
dramatically. The overall increase in number and size of fishing vessels indicate the
development of the economic importance of the crab. After 2001 the overall harvest
rate increased along with the growth of the stock.
The increase in crab stocks in recent years has resulted in severe by-catch issues,
particularly in the cod gillnet fishery. However some available size distribution data
for crabs caught by the gillnet fishery show that few juvenile specimens are caught.
Most crabs seem to be larger than CL 120 mm. More than 60% of the crabs caught
in the gillnet fishery in Varangerfjord were females, while large males dominate the
by-catch in the lumpsucker gillnet fishery during early summer. The by-catch of crabs
increased from 1997 to 1999, but declined in 2000–2002, and the estimated number
531The Invasive History, Impact and Management of the Red King Crab
Table 2 The number of vessels, fishing effort in traps allowed per boat, TAC, and size of the
vessels participating in the research- and commercial fishery of the red king crab in Norwegian
waters from 1994 to 2007 (From Hjelset et al. 2009)
of vessels
Fishing effort
traps per boat
TAC (legal
rate (%)
Overall vessel
Research fishery
1994 4 20 11,000 41 7–15
1995 4 20 11,000 11 7–15
1996 6 20 15,000 17 7–15
1997 6 20 15,000 14 7–15
1998 15 20 25,000 17 7–15
1999 24 20 38,000
2000 33 20 38,000 6 7–15
2001 116 20 100,000 22 7–15
Commercial fishery
2002 127 30 100,000 13 7–15
2003 197 30 200,000 15 7–15
2004 260 30 280,000 21 6–21
2005 273 30 280,000 34 6–21
2006 264 30 300,000 29 6–21
2007 253 30 300,000 31 6–21
Not available
in 2002 was a third as large as in 1999 (Sundet and Hjelset 2002; Hjelset et al. 2003).
This is probably due to the decline in the cod gillnet fishery. Low abundance of cod
has forced the fishermen to move further west along the coastline in search of fish,
thereby reducing the by-catch of the crab. The crab impacts the longline fishery by
removing the bait off the hooks, thereby reducing catches of target fish.
In order to compensate the fishermen for the loss of the traditional fishery
and equipment (i.e., gillnets, long-lines) caused by the invasion of the crab, the
criteria for participation in the annual fishery are set in favour of the local fish-
ermen. This is generally acknowledged by fishermen from other parts of
Norway, since the presence of the crab directly impacts the local fishermen
(Jørgensen et al. 2004).
7 Management and Future Challenges
From 1994 to 2001, the newly introduced red king crab stock was exploited through
a research fishery limited by TAC numbers (Table 2) in the territorial waters of
Russia and Norway. The harvest rate of the crab was relatively low (Sundet and
Hjelset 2002). Thereafter the management regime and the following harvest pattern
ensured that the largest males were removed from the population (Nilssen and
Sundet 2006).
532 L.L. Jørgensen and E.M. Nilssen
In 2002, the fishery had become commercial, and the Norwegian quota was set
at 100,000 crabs (Nilssen and Sundet 2006), and increased to 300,000 crabs in 2006
(Table 2). The management of the fishery was based on annual joint agreements
between Russia and Norway through the Mixed Russian-Norwegian Fishery
Commission. During 2004, Norway and Russia agreed to limit the spread of the
crab westwards by establishing a border at 26°E in the Norwegian zone (Fig. 3
North Cape). West of this longitude Norway was given free rein to apply all neces-
sary management methods with a view to limit the spread of the crab. The joint
Norwegian and Russian management ended in 2007. Since then management has
been continued by each country within their respective fishery zones in the Barents
At present two management regimes are implemented in Norwegian waters and
located to two different geographical areas/regions. One commercial eastern area
from the Russian border at 31°E to North Cape at 26°E which are controlled by the
governmental management plan for a king crab fishery where the population of
king crabs are managed in order to give the best possible biological and economical
output. The second area is the western area, south and west of 26° E, with a free
fishing of the red king crab in order to reduce the rate of spreading south along the
Norwegian coastline (St. meld. 40 2006; Øseth 2008).
The commercial stock in the eastern area is managed according to the ‘3-S’
regime (sex, size and season) and only males with a CL > 137 mm may be landed
(Nilssen and Sundet 2006). This strategy is similar to the Alaskan management
model (Otto 1986; Kruse 1993). In the western “free fishing area” all crabs are
landed without regard to size and sex.
It was not legal to land females CL > 137 mm in the eastern commercial manage-
ment area before 2008, but now allowed. This regime with an eastern commercial
managed area and a western free fishing area is still under evaluation and king crab
assessment and management in relation to harvest strategies, by-catch problems,
changes in gear technology, targeting ground fish and reducing the spread of this
invasive species is still under consideration (Jørgensen et al. 2007).
Both extended periods of heavy fishing pressure (Pollock 1995; Jørgensen et al.
2007) and lack of food can affect the life history traits of crustaceans. There will
always be a trade off between food available and the investment in growth, size/age
at maturation and reproductive output (Stearns 1992). Reduction in reproductive
output could be effected by lack of food which will be a consequence of the increased
biomass of crab. It is therefore necessary to investigate the variation in size at sexual
maturity and reproductive output in the population along the Norwegian coast in
order to establish a baseline for future management and monitoring (Hjelset et al.
2009). Therefore, registration of size at sexual maturity, fecundity and moulting
frequencies of the crab has been collected since 1992 and will be published in near-
est future.
Precise scientific predictions cannot be given concerning the future impacts of
the red king crab in the Southern Barents Sea. All indications suggest that this
invasive species will spread further north in the Barents Sea, as well as south-
wards along the coast of Norway. The possibility of transporting larvae in ballast
533The Invasive History, Impact and Management of the Red King Crab
water to other regions is an alarming reality, especially as the traffic of oil and gas
vessels around the Barents Sea and northern Norway is likely to increase in the
near future.
Acknowledgements Thanks to Geir Randby, Lillehammer Film and to Trond Thangstad for
illustrative figures. Thank to the editors of the volume and referees for language improvement and
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... aquaculture, ballast water), (ii) the implementation of early warning systems, and (iii) the application of proper RAs. By way of example, the introduction of the red king crab Paralithodes camtschaticus (Jørgensen & Nilssen, 2011) in the Barents Sea area during the 1960s and 1970s for commercial fishery enhancement is a remarkable case of mismanagement due to lack of implementation of proper RA, with the species' ecological and economic impacts having since been reported in several studies (Jørgensen & Nilssen, 2011). Implementing proper RA procedures for the management of non-native potentially invasive species is therefore crucial. ...
... aquaculture, ballast water), (ii) the implementation of early warning systems, and (iii) the application of proper RAs. By way of example, the introduction of the red king crab Paralithodes camtschaticus (Jørgensen & Nilssen, 2011) in the Barents Sea area during the 1960s and 1970s for commercial fishery enhancement is a remarkable case of mismanagement due to lack of implementation of proper RA, with the species' ecological and economic impacts having since been reported in several studies (Jørgensen & Nilssen, 2011). Implementing proper RA procedures for the management of non-native potentially invasive species is therefore crucial. ...
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Non-native marine crustaceans can exert detrimental impacts on native marine communities by altering habitat and ecosystem function. The Mediterranean Sea is particularly vulnerable to introductions of non-native crustaceans, as evidenced by their remarkably high establishment success. In this study, 20 species of non-native marine crustacean decapods and barnacles of which eleven extant and nine ‘horizon’ were screened for their potential invasiveness in the Mediterranean Sea. Using the Aquatic Species Invasiveness Screening Kit and including an additional nine native species to increase accuracy, calibrated risk thresholds of 3.5 for the BRA (Basic Risk Assessment) and 8.5 for the BRA+CCA (BRA + Climate Change Assessment) were obtained that distinguished reliably between invasive and non-invasive species. All 20 non-native species were classified as carrying a high risk of invasiveness for the Mediterranean Sea, both for the BRA and the BRA+CCA. Chinese mitten crab Eriocheir sinensis was by far the highest risk species, followed by Harris mud crab Rhithropanopeus harrisii, Asian shore crab Hemigrapsus sanguineus, Amphibalanus improvisus, and lesser swimming crab Charybdis (Goniohellenus) longicollis. The findings of this study will provide management and control directions for non-native marine crustaceans in the Mediterranean Sea, with special emphasis on regulations regarding ballast waters, which represent one of the main introduction pathways for these aquatic organisms.
... It has been reported that alien species can influence local habitats either by impacting native key species or by replacing them (Galil 2007); hence, such species can be ecologically and/or economically harmful (Williamson and Fitter 1996). This has been the case for three invasive crab species: the green crab Carcinus maenas in Tasmania (Walton et al. 2002), the Chinese mitten crab Eriocheir sinensis in San Francisco Bay (Rudnick et al. 2003), and the red king crab Paralithodes camtschaticus in the Barents Sea (Jørgensen and Nilssen 2011). In the case of P. segnis, Introduction published data are insufficient to assess its impact on Mediterranean benthic fauna. ...
... For example, the King crab (Paralithodes camtschaticus) has expanded westwards from the Barents Sea to the northeastern Norwegian coast (Jørgensen & Nilssen 2011). ...
Environmental stressors related to climate change and other anthropogenic activities are impacting Arctic marine ecosystems at exceptional rates. Within this context, predicting future scenarios of deep‐sea ecosystems and their consequences linked with the fate of coastal areas is a growing need and challenge. We used an existing food‐web model developed to represent the outer basin of the Malangen fjord, a Northern Norwegian deep‐sea ecosystem, to assess the potential effects of plausible future trajectories of change for major drivers in the area, including links to coastal kelp forests. We considered four major drivers (kelp particulate organic matter [POM] production entering the deep sea, fishing effort, king crab invasion, and ocean warming) to project 12 future scenarios using the temporal dynamic module of Ecopath with Ecosim approach. Overall, we found that the impact of warming on the deep‐sea ecosystem structure and functioning, as well as on ecosystem services, are predicted to be greater than changes in kelp forest dynamics and their POM production entering the deep‐sea and the king crab invasion. Yet, the cumulative impacts are predicted to be more important than non‐cumulative since some stressors acted synergistically. These results illustrate the vulnerability of sub‐Arctic and Arctic marine ecosystems to climate change and consequently call for conservation, restoration, and adaptation measures in deep‐sea and adjacent ecosystems. Results also highlight the importance of considering additional stressors affecting deep‐sea communities to predict cumulative impacts in an ecosystem‐based management and global change context and the interlinkages between coastal and deep‐sea environments. This article is protected by copyright. All rights reserved.
... The former is thought to have been introduced by ballast water, while the latter was an intentional introduction (Alvsvåg, Agnalt, & Jørstad, 2009;Jørgensen & Nilssen, 2011). The Barents Sea appears to be in transition from a cold Arctic to warm Atlantic climate regime (Lind, Ingvaldsen, & Furevik, 2018), making it particularly vulnerable to invasion. ...
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The risk of aquatic invasions in the Arctic is expected to increase with climate warming, greater shipping activity, and resource exploitation in the region. Planktonic and benthic marine aquatic invasive species (AIS) with the greatest potential for invasion and impact in the Canadian Arctic were identified and the 23 riskiest species were modelled to predict their potential spatial distributions at pan‐Arctic and global scales. Modelling was conducted under present environmental conditions and two intermediate future (2050 and 2100) global warming scenarios. Invasion hotspots – regions of the Arctic where habitat is predicted to be suitable for a high number of potential AIS – were located in Hudson Bay, Northern Grand Banks/Labrador, Chukchi/Eastern Bering seas, and Barents/White seas, suggesting that these regions could be more vulnerable to invasions. Globally, both benthic and planktonic organisms showed a future poleward shift in suitable habitat. At a pan‐Arctic scale, all organisms showed suitable habitat gains under future conditions. However, at the global scale, habitat loss was predicted in more tropical regions for some taxa, particularly most planktonic species. Results from the present study can help prioritize management efforts in the face of climate change in the Arctic marine ecosystem. Moreover, this particular approach provides information to identify present and future high‐risk areas for AIS in response to global warming.
... As the juvenile red king crabs most commonly are distributed in shallower and more sheltered areas (Pavlova 2008, Jørgensen & Nilssen 2011 than those used as spawning sites for the Barents Sea capelin (Gjøsaeter 1998), the predation on capelin eggs by juvenile red king crab is probably less important for capelin than for lumpsucker. When fitting the logistic regression model with the aim to predict the occurrence of capelin eggs in red king crab stomachs, the effect of crab size was insignificant (Paper III). ...
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Mortality of demersal fish eggs caused by predation was investigated in this research project. The selected prey species were the Barents Sea capelin Mallotus villosus and lumpsucker Cyclopterus lumpus L, while the selected predators were capelin itself by cannibalistic behaviour and the invasive red king crab Paralithodes camtschaticus. Both capelin and lumpsucker are commercially important species in the Norwegian fisheries, but capelin is also a very important forage fish for other species. The egg guarding lumpsucker male was not able to protect his eggs from the egg feeding red king crab. Approximately 8% of the analysed red king crab stomachs contained lumpsucker eggs in 2003. In 2005 and 2006, capelin eggs occurred in 10% and 23% of the analysed red king crab stomachs and capelin in 82% and 22% respectively. As post-spawn capelin occurred more frequently in crab stomachs than capelin eggs, they might have served as alternative prey to capelin eggs. Average stomach evacuation time for lumpsucker in red king crabs was higher than for capelin eggs. Consumption estimates of capelin eggs in red king crab accounted for 0.04% and 2.23% of the eggs available in the years of study, while the minimum estimated mortality due to egg cannibalism in capelin accounted for 1-2% of the total egg production in 2003. The uncertainty in stomach data generated most of the uncertainty in the Monte Carlo estimated consumption of fish eggs by the red king crab. This study has established new knowledge about fish egg predation by the invasive red king and egg cannibalism in capelin. Mortality in capelin eggs caused by cannibalism and egg consumption by the red king crab may influence mortality of eggs, but is not considered to hamper capelin recruitment. Recruitment of lumpsucker on the other hand, may be hampered by the red king crab that chase away the egg guarding male, damage and feed on his eggs
... While the introduction of the crab has brought economic benefits to Russia and Norway, there are concerns regarding its impacts on native communities, especially in Norwegian waters(Dvoretsky & Dvoretsky, 2015;Falk-Petersen, Renaud, & Anisimova, 2011;Oug, Cochrane, Sundet, Norling, & Nilsson 2011). To balance the economic benefits and ecological concerns, the Norwegian government implements two management regimes-a quota-regulated zone to sustain the crab population for exploitation and a free-fishing zone to reduce the rate of spread southward along the Norwegian coast(Jørgensen & Nilssen, 2011;Lorentzen et al., 2018). This example also highlights the need to coordinate NIS management strategies among Arctic nations, as introduced species may unintentionally spread to neighboring countries.Early detection of undesired NIS at potential high-risk regions, such as the Iceland Shelf, the Norwegian Sea, and the Barents Sea, is essential for protecting the Arctic region from new invasions.However, NIS may be overlooked if they are rare or morphologically cryptic, and sampling methods can result in false negatives (i.e., failure to detect the occurrence of NIS in a given environment) at the initial stage of an invasion(Delaney & Leung, 2010;Stanislawczyk, Johansson, & MacIsaac, 2018). ...
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Climate change and increased anthropogenic activities are expected to elevate the potential of introducing nonindigenous species (NIS) into the Arctic. Yet, the knowledge base needed to identify gaps and priorities for NIS research and management is limited. Here, we reviewed primary introduction events to each ecoregion of the marine Arctic realm to identify temporal and spatial patterns, likely source regions of NIS, and the putative introduction pathways. We included 54 introduction events representing 34 unique NIS. The rate of NIS discovery ranged from zero to four species per year between 1960 and 2015. The Iceland Shelf had the greatest number of introduction events (n = 14), followed by the Barents Sea (n = 11), and the Norwegian Sea (n = 11). Sixteen of the 54 introduction records had no known origins. The majority of those with known source regions were attributed to the Northeast Atlantic and the Northwest Pacific, 19 and 14 records, respectively. Some introduction events were attributed to multiple possible pathways. For these introductions, vessels transferred the greatest number of aquatic NIS (39%) to the Arctic, followed by natural spread (30%) and aquaculture activities (25%). Similar trends were found for introductions attributed to a single pathway. The phyla Arthropoda and Ochrophyta had the highest number of recorded introduction events, with 19 and 12 records, respectively. Recommendations including vector management, horizon scanning, early detection, rapid response, and a pan‐Arctic biodiversity inventory are considered in this paper. Our study provides a comprehensive record of primary introductions of NIS for marine environments in the circumpolar Arctic and identifies knowledge gaps and opportunities for NIS research and management. Ecosystems worldwide will face dramatic changes in the coming decades due to global change. Our findings contribute to the knowledge base needed to address two aspects of global change—invasive species and climate change.
... Other documented ecological impacts of this decapod include reduced density and biomass of benthic invertebrates (Britayev et al. 2010), a shift in size composition towards smaller, motile organisms, effects on sediment quality (Oug et al. 2011) and predation on demersal fish eggs (Mikkelsen & Pedersen 2012). Despite a considerable interest in understanding the impact of the red king crab on the benthic ecosystem (Gudimov et al. 2003, Jørgensen & Primicerio 2007, Jørgensen & Nilssen 2011, Oug et al. 2011, indirect food web effects (e.g. competition) remain poorly studied, and overall impacts on the ecosystem structure and stability are unknown. ...
Since the 1990s, the density of the invasive red king crab Paralithodes camtschaticus has increased dramatically in coastal areas in northern Norway. We investigated its direct and indirect effects on food web structure and ecosystem properties (e.g. species biomasses and production) in the Porsanger Fjord in the study period 2009−2011 using 5 subarea Ecopath food web models. The 5 baseline models with different red king crab densities were compared and the food web effects of crab removal were explored through simulations in Ecosim. King crabs were important as benthic predators and exerted strong top-down effects on long-living invertebrates such as predatory gastropods, asteroids, detritivorous echinoderms and herbivorous sea urchins. The crab experienced little predation from fish or other predators at higher trophic levels, thus food web effects of the red king crab generally stayed within the benthic compartment. Red king crab removal decreased system omnivory and resulted in higher food web biomass−low turnover systems, with relatively lower production:biomass ratios of benthic invertebrates. Other ecosystem properties (e.g. total production, consumption, ascendancy and overhead) were little affected by crab abundance and suggest stable systems. Effects of crab removal were less significant in baseline models with low initial crab biomass, and high benthic production by detritivores in the inner fjord may buffer future predation in this area. Indirect effects of crab predation included a positive cascade effect on macroalgae due to predation on herbivorous sea urchins and a negative effect on benthic-feeding birds, indicating competition for invertebrate prey.
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The red king crab (RKC, Paralithodes camtschaticus) is a highly-valued decapod species. Typically, RKCs undergo a period of live holding (LH), often without feeding, in onshore facilities, allowing for flexible management before export to destination markets. This study aimed to (i) gain information on the fatty acid (FA) profile of the cooked leg meat and raw hepatopancreas obtained from RKC harvested in Norwegian waters of the Barents Sea and (ii) investigate how these FA profiles are affected by LH without feeding for up to 92 days at 5 or 10 °C. Minor changes were observed in the FA profile of cooked leg meat, which retained its nutritional value in omega−3 FA content. In contrast, the FA composition of raw hepatopancreas was severely affected by the LH time, with substantial changes occurring especially between 41 and 62 days at 10 °C and between 62 and 92 days at 5 °C. Saturated and specific monounsaturated FAs, such as 16:1n−7c and 18:1n−9t, as well as 22:5n−3c, were preferentially utilized at the beginning of the starvation period, followed by the mobilization of C18–22 unsaturated FAs. Long-chain highly-unsaturated FAs were preferentially retained during LH, especially 20:4n−6c and 22:5n−3c. The information emerging from the present study may be practically exploited for selecting or designing suitable feed for RKC during LH at different temperatures.
The Barents Sea Region differs from most other Arctic sea areas in the way that the majority of the Barents Sea has historically been ice-free all year round. There is a huge human population living around the Barents Sea, and the exploration of both living and non-living resources is important. The region is highly developed and industrialized. Cold Arctic waters meet warm Atlantic waters and there are diverse and different ecosystems between the northern and southern part of the region. Due to climate change and warmer waters there is an ongoing borealization of the Barents Sea region. Species are moving northwards and new species are introduced to the area. The region is one of the most fishery intensive regions in the world. Because of the warmer waters, the huge northeast Arctic cod population is moving farther north and east in the Barents Sea. The commercially important species mackerel and snow crab have migrated to the region. The co-management regime between Norway and Russia is a main reason for the healthy fish stocks in the region.
Technical Report
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Diagnóstico de las pesquerías argentinas. Informe elaborado en el marco de la etapa de preparación del proyecto de inversión GEF/FAO "Fortalecimiento de la la Gestión y Protección de la Biodiversidad Costero Marina en áreas ecológicas clave y la aplicación del enfoque ecosistémico de la pesca (EEP)" Introducción 5 Contexto General 5 Descripción y composición de la flota 8 La flota pesquera nacional 9 La estadística pesquera 10 Marco Institucional 13 Metodología 16 CARACTERÍSTICAS DE LAS UNIDADES ECOSISTÉMICAS 23 Unidad Ecosistémica I: Frente Marítimo Argentino Uruguayo – somero Bonaerense 23 1. Corvina rubia Micropogonias furnieri 23 2. Pescadilla de red - Cynoscion guatucupa 29 3. Variado costero bonaerense - Condrictios 33 Unidad Ecosistémica II: Bonaerense – El Rincón y plataforma media 37 4. Caballa Scomber japonicus 37 Variado costero bonaerense - Condrictios 40 5. Anchoita Engraulis anchoita 40 Merluza común - Merluccius hubbsi 46 Unidad ecosistémica III: Patagonia norte Golfo San Jorge – Península de Valdés 47 6. Merluza común - Merluccius hubbsi 47 7. Abadejo Genypterus blacodes 56 8. Langostino– Pleoticus muelleri 59 9. Calamar - Illex argentinus 65 10. Centolla - Lithodes santolla 70 Unidad Ecosistémica IV: Talud 73 Ecosistema IVa: Talud norte 73 Merluza común – Merluccius hubbsi 73 Calamar - Illex argentinus 73 11. Vieira patagónica - Zygochlamys patagonica 73 Unidad ecosistémica IVb: Talud sur 79 Calamar - Illex argentinus 79 12. Polaca Micromesistius australis 79 13. Merluza de cola - Macruronus magellanicus 83 Ecosistema V: Sistemas costeros de golfos norpatagónicos: Golfo San Matías, Golfo San José y Golfo Nuevo 87 14. Merluza común - Merluccius hubbsi 89 15. Savorín Seriolella porosa 89 16. Pesquerías bentónicas (mejillón, vieira tehuelche, almeja púrpura, pulpito patagónico y cangrejo nadador) 90 16.a Mejillón Mytilus edulis platensis 90 16.b Vieira tehuelche Aequipecten tehuelchus 90 16.c Almeja púrpura Amiantis purpurata 92 16.d Pulpito patagónico Octopus tehuelchus 92 16.e Cangrejo nadador Ovalipes trimaculatus 93 Otros 93 Ecosistema VI: Plataforma austral 94 Polaca Micromesistius australis 94 Merluza de cola - Macruronus magellanicus 94 Literatura citada 95 Anexo I 112 Anexo II 114
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The red king crab (Paralithodes camtschaticus) was intentionally transferred from areas in the Northern Pacific Ocean to the Russian Barents Sea during the 1960s (1961–1969), to create a new and valuable commercial resource. A reproductive population in the receptor region was evident ten years later and from this time the species has continued to spread both north and east in the Barents Sea and southwards along the coast of Northern Norway. Ecological im-pacts upon the native fauna are investigated through, among others, analysis of the diet of the crab, as molluscs, echinoderms, polychaetes and crustaceans are frequently found as prey items. Problems following the invasion of the red king crab are displayed as bycatch of crabs in gill-net- and longline-fisheries. The crab is regarded as a commercial resource both in Russia and Norway. Management of the red king crab is undertaken as a joint stock between Norway and Russia through the Joint Russian-Norwegian Fishery Commission
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The relationships between growth rates, fecundity and lengths and ages at sexual maturity are explored, with examples primarily in spiny lobsters, to illustrate the underlying principles involved. Changes in the dynamics of exploited populations of both fish and crustaceans often have important implications for egg production. Two independent processes are thought to be involved, genetic changes caused by the removal of certain genotypes by size-selective fisheries, and growth changes caused by altered stock densities and/or changed food availability. The impacts of these processes on maturity length appear to be antagonistic, giving rise to some contradictory effects in different exploited stocks, some stocks exhibiting increases and others showing decreases in length at maturity.
SYNOPSIS. Non-indigenous species (NIS) are increasingly conspicuous in marine and estuarine habitats throughout the world, as the number, variety, and effects of these species continue to accrue. Most of these NIS invasions result from anthropogenic dispersal. Although the relative importance of different dispersal mechanisms varies both spatially and temporally, the global movement of ballast water by ships appears to be the largest single vector for NIS transfer today, and many recent invasions have resulted from this transfer. The rate of new invasions ' may have increased in recent decades, perhaps due to changes in ballast water transport. Estuaries have been especially common sites of invasions, accumulating from tens to hundreds of NIS per estuary that include most major taxonomic and trophic groups. We now know of approximately 400 NIS along the Pacific, Atlantic and Gulf coasts of the U.S., and hundreds of marine and estuarine NIS are reported from other regions of the world. Although available information about invasions is limited to a few regions and underestimates the actual number of NIS invasions, there are apparent differences in the frequency of NIS among sites. Mechanisms responsible for observed patterns among sites likely include variation in supply of NIS, and perhaps variation in properties of recipient or donor communities, but the role of these mechanisms has not been tested. Although our present knowledge about the extent, patterns and mechanisms of marine invasions is still in its infancy, it is clear that NIS are a significant force of change in marine and especially estuarine communities globally. Taxonomically diverse NIS are having significant effects on many, if not most, estuaries that fundamentally alter population, community, and ecosystems processes. The impacts of most NIS remain unknown, and the predictability of their direct and indirect effects remains uncertain. Nonetheless, based upon the documented extent of NIS invasions and scope of then effects, studies of marine communities that do not include NIS are increasingly incomplete.
Feeding and growth in wet weight of the red king crab Paralithodes camtschaticus were studied under laboratory conditions for 4 months from September 1993 to January 1994. Crabs were divided into 3 groups: ovigerous females, juvenile females, and mature males. Food consumption (g · d-1) significantly increased with crab wet weight (W), while feeding rate (FR, g · kg-1 · d-1), weight-standardized food consumption, decreased significantly with crab weight. The relationship of FR ~ W significantly differed between ovigerous females and juvenile females and males, but not between juvenile females and males, and can be expressed as FR = 70.9 - 0.017 W (P < 0.001) for ovigerous females, FR = 64.1 - 0.017 W (P < 0.001) for juvenile females and males. Crabs ceased feeding during molting, and feeding rates were significantly lower between 12 days before and 8 days after ecdysis than during the nonmolting period. Molted male crabs rapidly increased weight after molt. Their growth rate averaged 412.3 g · kg-1 ± 89.6 SD (wet weight) and decreased with crab weight. Among unmolted crabs, growth rates significantly differed between ovigerous females (66.6 g · kg-1 ± 19.4 SD), juvenile females (50.4 g · kg-1 ± 14.5 SD), and males (5.2 g · kg-1 ± 4.0 SD).
ABSTRACT Seasonal movements and distribution of primiparous and multiparous red king crabs (Paralithodes camtschaticus) were monitored with ultrasonic biotelemetry approximately weekly for 1 year in Auke Bay, Alaska. Migration was associated with life-history events and may have occurred in response to spatial and temporal variations in environmental conditions and resources. All crabs displayed distinct shifts in depth and habitat use and followed a general pattern of seasonal movement as follows: (1) gradual movement to deep water in spring after mating and egg extrusion, and residence there through early November; (2) abrupt, synchronous movement into shallow-water areas in November, and residence there through late February or early March; and (3) gradual, synchronous movement to intermediate depths followed by movement into shallow water to molt and mate between late March and late May. The behavior of primiparous crabs was more variable than that of multiparous crabs. The differences in behavior may result from ontogenetic shifts in movements, and habitat selection. The annual range of primiparous crabs (x = 11.9 km2) exceeded that of multiparous crabs x( = 3.6 km2). Mean depth was directly correlated with photoperiod, and the sudden, synchronous movement of crabs between habitats coincided with thermohaline mixing. Females displayed a highly aggregated distribution, especially during winter in shallow water, where podding behavior of adult crabs was documented for the first time.