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521
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-mail: lis.lindal.joergensen@imr.no
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)
Year
Varangerfjorden
CPUE ± CI
Tanafjorden
CPUE ± CI
Laksefjorden
CPUE ± CI
1994
a
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
a
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
populations.
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)
Year
Number
of vessels
Fishing effort
traps per boat
TAC (legal
males)
Harvest
rate (%)
Overall vessel
length(m)
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
a
7–15
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
a
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
Sea.
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
comments.
References
Anisimova N, Berenboim B, Gerasimova O, Manushin I, Pinchukov M (2005) On the effect of red
king crab on some components of the Barents Sea ecosystem. Ecosystem dynamics and opti-
mal long-term harvest in the Barents Sea Fisheries. Proceeding of the 11th Russian-Norwegian
Symposium, Murmansk, 15–17 Aug 2005/ IMR/PINRO Joint Report Series 2:298–306
Bright DB (1967) Life histories of the king crab, Paralithodes camtschatica, and the Tanner crab,
Chionoecetes bairdi, in Cook Inlet, Alaska. PhD thesis, University of South California, Los
Angeles
Carlton JT (1996) Pattern, process, and prediction in marine invasion ecology. Biol Cons
78:97–106
Cunningham DT (1969) A study of the food and feeding relationships of the Alaskan king crab
Paralithodes camtschatica. MSc thesis, State College, California, San Diego
Dew CB (1990) Behavioural ecology of podding red king crab, Paralithodes camtschatica. Can J
Fish Aquat Sci 47:1944–1958
Elton CS (1958) The ecology of invasions by animals and plants. Methuen, London
Faccia I, Alyakrinsky A, Bianchi CN (2009) The crab that came in from the cold: first record of
Paralithodes camtschaticus (Tilesius, 1815) in the Mediterranean Sea. Aquat Invas 4:715–718
Fenyuk VF (1945) Analiz soderzhimogo zheludkov Kamchatskogo craba [Analysis of stomach
contents of the Kamchatka crab]. In: Materialy po biologii, promyslu i obrabotke Kamchatskogo
kraba [Materials on biology, fishery and refinement of the Kamchatka crab]. Izv tikhookean
nauchnoIssled Inst ryb khoz Okeanogr 19:71–78 (In English, Japanese, Russian)
Fukuhara FM (1985) Biology and fishery of south-eastern Bering Sea red king crab (Paralithodes
camtschatica, Tilesius). NOAA Proc Rep 85-11:801-982
Gerasimova OV (1997) Analysis of king crab (Paralithodes camtscahtica) trophic links in the
Barents Sea. ICES CM 1977/GG:03
Gudimov AV, Gudimova EN, Pavlova LV (2003) Effect of the red king crab Paralithodes
camtschaticus on the Murmansk coastal macrobenthos: the first estimates using sea urchins of
the genus Strongylocentrotus as an example. Dokl Biol Sci 393:539–541
Hansen T (2002) Temperaturpreferanser hos kongekrabbe (Paralithodes camtschaticus)
[Temparature preferences of the red king crab (Paralithodes camtschaticus)]. Cand. scient.
Thesis in Marine Biology, Norwegian College of Fishery Science, University of Tromsø,
Norway, pp 86 [In Norwegian] http://www.angelfire.com/on/2mash/Filer/hovedfaget.pdf
Haugan TA (2004) Bunnsamfunn og næringsvalg hos kongekrabbe, Paralithodes camtscahaticus
(Tilesius, 1815), på noen lokaliter i Finnmark. (Benthic community an food preferences by
king crab Paralithodes camtschaticus (Tilesius, 1815), at some localities in Finnmark). MSc
thesis, Norwegian College of Fishery Science, University of Tromsø, Norway (In
Norwegian)
Herrick FH (1909) Natural history of the American lobster. Bur Fish Bull 29:149–408
Hjelset AM, Pinchukov MA, Sundet JH (2003) Joint report for 2003 on the red king crab
(Paralithodes camtschaticus) investigations in the Barents Sea. Report to the 32 h Session for
the Mixed Russian-Norwegian Fisheries Commission, pp 13
534 L.L. Jørgensen and E.M. Nilssen
Hjelset AM, Sundet JH, Nilssen EM (2009) Size at sexual maturity in the female red king
crab (Paralithodes camtschaticus) in a newly settled population in the Barents Sea, Norway.
J Nortw Atl Fish Sci 41:173–182
Jewett SC, Feder HM (1982) Food and feeding habits of the king crab Paralithodes camtschatica
near Kodiak Island, Alaska. Mar Biol 66:243–250
Jewett SC, Onuf CP (1988) Habitat suitability index models: red king crab. US Fish Wildl Ser Biol
Rep, National Wetlands Research Center, Slidell Louisiana 82: 34
Jewett SC, Gardner LA, Rusanowski PM (1989) Food and feeding habits of red king crab from
north-western Norton Sound Alaska. In: Proceedings of the International Symposium on King
Tanner Crabs. University of Alaska Sea Grant Report 90–04:219–232
Jørgensen LL (2005) Impact scenario for an introduced decapod on Arctic epibenthic communi-
ties. Biol Invas 7:949–957
Jørgensen LL, Primicerio R (2007) Impact scenario for the invasive red king crab Paralithodes
camtschaticus (Tilesius, 1815) (Reptantia, Lithodidae) on Norwegian, native, epibenthic prey.
Hydrobiologia 590:47–54
Jørgensen LL, Manushin I, Sundet JH, Birkely S-R (2004) The intentional introduction of the
marine Red King Crab Paralithodes camtschaticus into the Southern Barents Sea, pp 124–137.
In: ICES (ed) Report of the Working Group on Introductions and Transfers of Marine
Organisms (WGITMO). ICES Special Advanced Report Cesenatico, Italy
Jørgensen C, Enberg K, Dunlop ES, Arlinghaus R, Boukal DS, Brander K, Ernande B, Gårdmark A,
Johnston F, Matsumura S, Pardoe H, Raab K, Silva A, Vainikka A, Dieckmann U, Heino M,
Rijnsdorp AD (2007) Managing evolving fish stocks. Science NY 318:1247–1248. doi:10.1126/
science.1148089
Kruse GH (1993) Biological perspectives on crab management in Alaska. In: Kruse GH, Eggers
DM, Marasco RJ, Pautzke C, Quinn II TJ (eds) Proceedings of the international symposium
on management strategies for exploited fish populations. University of Alaska Sea Grant
College Progress Report 93-02:355–384
Kulichkova VA (1955) Pitanie Kamchatskogo kraba v vesenneletnii period u beregov Kamchatki
i Sakhalina [The feeding pattern of the Kamchatka crabs off the coasts off Kamchatka and
Sakhalin]. Izv tikhookean nauchno Issled Inst ryb khoz Okeanogr 43:21–42 (In Russian)
Kun MS, Mikulich LV(1954) Sostav pishchi dalnevostochnykh promy slovikh krabov v letnii
period [Diet composition of Far Eastern crabs of commercial quality during the summer]. Izv
tikhookean nauchno Issled Inst ryb khoz Okeanogr 41:319–332 (In Russian, Japanese and
English)
Kurata H (1961) On the age and growth of king crab, Paralithodes camtschatica. Hokkaido Fish
Exp Sta 18:10–22, Monthly Rep
Larsen L (1996) Temperaturafhængig udvikling, vækst og dødelighed hos larver av kongekrabben
(Paralithodes camtschatica Tilesius) under eksperimentelle forhold [Temperature depended
development, growth and mortality of Red king crab (Paralithodes camtschatica Tilesius)
larvae in experimental conditions]. Candidate scientific thesis in Marine Biology, Norwegian
College of Fishery Science, University of Tromsø, Norway, pp 86 (In Danish)
Lodge DM (1993) Biological invasions: lessons for ecology. Trends Ecol Evol 8:133–137
Logvinovich DN (1945) Akvarial’nya nablyudeniya nad pitaniem Kamchatskogo kraba [Aquarium
observations on the feeding of the Kamchatka crab]. In: Materialy po biologii promyslu o
obrabotke Kamchatskogo kraba [Materials on biology, fishery and refinement of the
Kamchatka crab]. Izv tikhookean nauchno Issled Inst ryb khoz Okeanogr 19:79–97 (In
Russian, Japanese, and English)
Marukawa H (1933) Taraba-gani chosa [Biological and fishery research on the Japanese king crab
Paralithodes camtschatica (Tilesius)]. J Imp Fish Exp Sta 4:152 (In Japanese with English
abstract)
Nilssen EM, Sundet JH (2006) The introduced species red king crab (Paralithodes camtschaticus)
in the Barents Sea II Growth increments and moulting probability. Fish Res 82:319–326
Orlov YI, Ivanov BG (1978) On the introduction of the Kamchatka king crab Paralithodes
camtschatica (Decapoda: Anomura: Lithodidae) into the Barents Sea. Mar Biol 48:373–375
535The Invasive History, Impact and Management of the Red King Crab
Orlov YI, Karpevich AF (1965) On the introduction of the commercial crab Paralithodes
camtschatica (Tilesius) into the Barents Sea. Rapp P-v Réun Cons Int Explor Mer 156:59–61
(In: Cole HA (ed) ICES Spec. Meeting 1962 to consider problems in the exploitation and regu-
lation of fisheries for Crustacea)
Øseth E (2008) Forvaltning av kongekrabbe (Paralithodes camtschaticus) – et økologisk eksperi-
ment? (Management of king crab Paralithodes camtschaticus – ecological experiment?).
MSc thesis, University of Tromsø (In Norwegian) http://henry.ub.uit.no/munin/bitstream/10037/
1377/1/Masteroppgaven.pdf
Otto RS (1986) Management and assessment of eastern Bering Sea king crab stocks. In: Jamieson
GS, Bourne N (eds) North Pacific Workshop on stock assessment and management of inverte-
brates. Can Spec Publ Fish Aquat Sci 92:83–106
Pavlova LV (2009) Estimation of foraging on the sea urchin (Strongylocentrotus droebachiensis
(Echinoidea: Echinoida) by the red king crab Paralithodes camtschaticus (Malacostraca:
Decapoda) in coastal waters of the Barents Sea. Russ J Mar Biol 35:288–295
Pavlova LV, Britayev TA, Rzhavsky AV (2007) Benthos elimination by juvenile red king crabs
Paralithodes camtschaticus (Tilesius, 1815) in the Barents Sea coastal zone: experimental
data. Dokl Biol Sci 414:231–234
Pedersen OP, Nilssen EM, Jørgensen LL, Slagstad D (2006) Advection of the Red king crab larvae
on the coast of North Norway – a Lagrangian model study. Fish Res 79:325–336
Pollock DE (1995) Changes in maturation ages and sizes in crustacean and fish populations. S Afr
J Mar Sci 15:99–103
Powell GC (1974) Gregarious king crabs. Sea Frontier 20:206–211
Powell GC, Nickerson RB (1965a) Aggregations among juvenile king crabs (Paralithodes
camtschatica, Tilesius) Kodiak, Alaska. Anim Behav 13(2–3):374–380
Powell GC, Nickerson RB (1965b) Reproduction of king crabs Paralithodes camtschatica
(Tilesius). J Fish Res Bd Can 22:101–111
Powell GC, Reynolds RE (1965) Movements of tagged king crabs Paralithodes camtschatica
(Tilesius) in the Kodiak Island - Lower Cook Inlet Region of Alaska, 1954–1963. Alaska Dep.
Fish Game Info Leafl 55, p 10
Rodin VE (1990) Population biology of the king crab Paralithodes camtschatica Tilesius in the
North Pacific Ocean. In: Proceedings of the International Symposium on King Tanner Crabs.
Univ Alaska Sea Grant Rep 90-04:133–144
Ross DJ, Johnson CR, Hewitt CL (2003) Variability in the impact of an introduced predator
(Asterias amurensis: Asteroidae) on soft-sediment assemblages. J Exp Mar Biol Ecol
288:257–278
Ruiz GM, Carlton JT, Grosholz ED, Hines AH (1997) Global invasion of marine and estuarine
habitats by non-indigenous species: mechanisms, extent, and consequences. Am Zool
37:621–632
Shirley SM, Shirley TC (1989) Depth and diel feeding periodicity of red king crab larvae. Am
Zool 29:64A, Abstract No. 280
St. meld. 40 (2006–2007) Forvaltning av kongekrabbe. (Management of the king crab). Ministry
Fisheries and Coastal Affairs (In Norwegian) http://www.regjeringen.no/en/dep/fkd/Documents/
Propositions-and-reports/stmeld/2006-2007/stmeld-nr-40-2006-2007-.html?id=480559
Stearns SC (1992) The evolution of life histories. Oxford University Press, Oxford, p 249
Stone RP, O’Clair CE, Shirley TC (1992) Seasonal migration and distribution of female red king
crabs in a Southeast Alaskan estuary. J Crust Biol 12:546–560
Sundet JH (2008) Bestandsvurdering av kongekrabbe i 2008 (Stock assessment of king crab in
2008). Intern Rep, pp 20 (In Norwegian)
Sundet JH, Hjelset AM (2002) The Norwegian red king crab (Paralithodes camtschaticus); man-
agement and by catch issues. In: Paul AJ, Dawe EG, Elner R, Jamieson GS, Kruse G H, Otto RS,
Sainte-Marie B, Shirley TC, Woodby D (eds), Crabs in Cold Water Regions: Biology,
Management, and Economics. University of Alaska Sea Grant Report AK-SG-02-01:681–692
Sundet JH, Rafter EE, Nilssen EM (2000) Stomach content of the red king crab (Paralithodes
camtchaticus) (Tilesius, 1815) in the southern Barents Sea. Crustacean issues 12. The biodiversity
536 L.L. Jørgensen and E.M. Nilssen
crisis and Crustacea. Proceedings of the forth international crustacean congress, Amsterdam,
Netherlands 20–24 July 1998, 2:193–201
Takeuchi I (1967) Food of king crab, Paralithodes camtschatica, of the west coast of Kamchatka
Peninsula, 1958–1964. Bull Hokkaido Reg Fish Res Lab 33:32–44 (In Japanese with English
summary) Also as: Fish Res Bd Can Transl. Ser., 1194 (1968)
Tsalkina AV (1969) Characteristics of the epifauna of the West Kamchatka shelf. Fish Res Bd Can
Transl Ser 1568
Vinogradov LG (1969) On the re production mechanism in the stock of the Kamchatka crab,
Paralithodes camtschatica, in the Okhotsk Sea off Western Kamchatka. Fis Res Bd Can Transl
Ser 1540
Wallace MM, Pertuit CJ, Hvatum AR (1949) Contribution to the biology of the king crab,
Paralithodes camtschatica Tilesius. US Dep Inter Fish Wildl Serv Fish Leafl 340:50
Walton WC, MacKinnon C, Rodriguez LF, Proctor C, Ruiz GM (2002) Effect of an invasive crab
upon a marine fishery: green crab, Carcinus maenas, predation upon a venerid clam, Katelysia
scalarina, in Tasmania (Australia). J Exp Mar Biol Ecol 272:171–189
Zelina DA, Mugu NS, Volkov AA, Sokolov VI (2008) Red king crab (Paralithodes camtschaticus)
in the Barentse Sea: a comparative study of introduced and native populations. Russ J Genet
44:859–866
Zhou S, Shirley TC, Kruse GH (1998) Feeding and growth of the red king crab Paralithodes
camtschaticus under laboratory conditions. J Crust Biol 18:337–345
