H)vfmJnologw 503: 21-2il. 2003,
M.8. Jon"!', A. {,,/Sol/slo", £ Olo.!<'«JI]. G l' fldgasrm, K. GUIII'W,...,,,, &,
J Smmruoll ("ds), Mi/:ri1tiOflS a"d Di.f{'I''7wl Q{MarilU': 0l1:dI,;sllrs.
02003 KI"...",. ,Imdt'''';c P"blishl'rs. Pnnll'd i" In" N",herIOllds.
21
Spread of the Chinese mitten crab (Eriocheir sinellsis H. Milne Edwards)
in Continental Europe: analysis of a historical data set
L-M. Herborg!, S. P. Rushton 2, A. S. Clare! & M. G. Bentley!
I School
ofMarine Science & ledlllO[ogy, Ridley Building, University of Newcastie-llpon-Tyne, Newcastle
upon TYI/e. NEl 7RU.
u.K.
2Cemre!or Life Science Modelling. Schoolo/Biolog;; Porler Building. VI/ire,s;l)' of NeIl'CaSlle-Il/JOII-T)'Ile,
Newcasrll' IIpon Tyne, NEI 7RU, U.K.
E-mail: leif- lIl atthias.herborg@llc/.llc.uk
56 4 2 0
Key lI'ords: invasive species. Eriocheir sinensis. crustacean, migration. catadromous. Europe
Abstract
The Chinese minen crab, Eriocheir sinensis (H. Milne Edwards). is an invasive species (hat lives as an adul!
predominantly in freshwater but migrates seawards to breed. II has spread 'ria ballasl water andlor intentional
introduction 10 Continental Europe, Southern France. U.S.A. (San Francisco Bay) and the United Kingdom.
An31ysis of detailed historic data from the outbreak in Europe was digitised and analysed using Geographical
Information Software. This revealed th3t Ihere were two separale invasions in Northern Europe and Sonthern
France. wilh an average range expansion during the peak period of 562 kmIyear from 1928-1938 (Nonhern Europe)
and 380 kmfye3r from 1954-1960 in Soulhern France. Size class distribution dflla from the lower estuary of the
River Elbe (Germany) (1932-1936) illustrate migration patterns to and from Ihe estuary oyer the year. Marking
experiments determined that lhe mean rate of downstream migration for adults was 11.5 kmJday (SO 3.54; n=7).
up to a maximum of 18.1 kmlday. The carapace width of upstream-migrating animals increased by 3 mmllOO
kill. The peak period for upstream migration was March 10 July, followed by the downstream season from July 10
September. This data set. cxtmclcd frOIll historic references. represents one of the most complete pictures of the
life cycle and spreading behaviour of this alien invader.
Introduction
The Chinese minen crab. Eriocheir sinensis (H. Milne
Edwards). originales in the Far &1St. its native range
extending from Hong Kong (.'~:~22° N) to the border
with North Korea ("':::40 0 N) (Hymanson et aI., 1999).
II is catadromous returning to Ihe sea 10 reproduce
once or rarely twice and then die. During the upstream
migralion of juveniles from estuaries, minen crabs can
reach rivers, lakes, and ponds as far as 1200 k1l1 from
the coasl (Peters, 1933). There they grow and mature.
The age at which E. sinensis reaches sexual maturity
varies from 1-3 years in China (Jin et a1.. 2001) to 3-5
years in Europe (Schubert. 1938). The Chinese millen
crab rcturns, when sexually mature at the end of its
life cycle, to estuaries for reproouclion in late summer.
After mating, females move into Ihe more marine pans
of the estuary and release their larvae in early spring.
Similar to E. ja"OIlica (de Haan), E. sinensis releases
up to three batche.~ of eggs (Peters. 1933; Kobayashi
& Matsuura. 1999: Kobayashi. 2001). Afler one reproductive season. the life cycle of most E. sinensil'
draws 10 an end and Ihcy dic (Peters, 1938c). Larval
developmcnt consists of 6 pelagic stages (I prezoea, 5
zoea) lasling ca. 90 days al 12 °C(Anger. 1991: Montu
et aI., 1996). After selliement; juvenile crabs migrate
upstream. The pelagic larval Slagcs arc thc JIlost likely
vector of spread for E. sinensis in ship's bal1ast water
(Peters. 1933; Cohen & Carhon, 1997).
Tbis invasive species has several impacts on humans and biodiversity. Firstly, it interferes with recreational and commercial fishing; in the San Francisco
Bay area. the presence of millen crabs has lead to
commercial shrimp trawlers abandoning certain fish-
22
1930
·-rJ
!.
.-
I
i
."
1960
,
.,'"
,-
)->;,:"~,,
Figfll"t
~'r
/
/
J. Cumulative sighlings of /.;"riQChtrr £it~1I!ijs from the literature across C()nlincntal !;uR')pe between 1912 and 1910.
ing areas. In the same area, there have been numerous
complaints from recrentiOl1al fishermen who blame the
crabs for causing an increased los:. of bait (Veldhuizen
& Swnish, 1999). Similar problems have been recorded In Europe where lnglc (19~6) reponed that, in
1981. Dutch fishennen suffered serious net damage
from mass occurrence of the mitten crab. Secondly. the
burrow-digging habit of E. sinensis can cause serious
river bank erosion. This is observed mainly in lidally-
influenced areas. or other stretches of ovcrs with
fluctuations in water level. Burrows are made in river
banks with a step gradiellt. and the necessary structural
strength to allow burrowing. They can be localed anywhere between the high-watcr murk, under recd banks
to below the low-water mark with the highest occurrencc around mid- water level (up to 301m 2) (Peters,
1933, 1938b; Dutton & Conroy, 1998). Thirdly, E. sinc/lSis may compete for resuurces with local freshwater
fish and invertebrates. Overlap of dietary and habitat
preferences with the economicallmpor1ant introduced
signal crayfish (Pacifastocus lefliltsculus, Dana) occurs in San Francisco Bay (Veldhuizen & Stanish.
1999). In Britain. the already threatened nativc crayfish (AlIstfOlwtamobiu$ pall'/Je.t. Lereboullel) could
w..- - - - - - - - - - - - - - - - - - - - - - - ' "
23
face similar prc.,sure (Clark el ai., 199~). Large numbers of millen crab:. occurring in some areas. their
large Si7C and aggressi ...c behaviour, in combination
with overlap in feeding and habitat selection. gives rise
to concerns about the possible impacts of current and
future invasions (Clark & Rainbow, 1996).
The sight of thousand.'> of crabs migrating over-
land around weirs in Europe in lhe 1930s was a major
(aclOr in raismg public awareness. TIle scale of the
occurreoce of Ihl" Cnl<;tace:m in rivers and lakes. especialJ ... during migrations has caused suffiCIent public
inte;est to engender many publications documenting
liS spread. Some of these pco\,jde much data on population distnbullon. migration patlems and spread particularly in Germany (Ilelers, 1933. 1938a: Panning,
1933.19380. b). The aim of this study was to analyse
the inva~lOn of E. sinensIS in comincntal Europe, using
literature conlempOr:lry with thl: major outbreak in.
small as 2 lllm) were collected ill large numbers from
the banks of the river Ellx: throllghotltthc year: larger
uabs were collceted on an occa~ional hasis from the
catches of fishermen using bOllom trawls. gill nelS and
eel buckets. Size classes within the population in the
Elbe were delermifll.:O ll<;.ing modal cla"s progression
unaly<:ls (Bhanacharya's method) In the FiSAT stock
assessment package. Using this method. different cohorts wilhin the monthly size-frequency distribution
dal;l could be identifi.. .d , and a mean and standard
denation derived. It should be nOled Ihat all SIZe measurements used in the historic liternture are in mm
carapace length (el). rather than the carapace width
used in morc reccllt literature. Therefore. carapace
length is used during the analysis: IhlS can be transformed into carapace \... idth using the formula derrived
by Peters (1933):
carapa....-c width (nun) = 1.13x Cilrdj);Jl:e length (mm).
MatHials and methods
TIle spread characlcristics of the Chlllc<;c mitten Cr:lb
during two invasive events m Nonhern Europe (ca.
1910-1950) and in Southem France (ca. 1950-19(0)
were quantified. TIlis was achieved by digitising all
slI;hlings (/,=646) fromlhe liter.lIure (Fig. I) and using GeographIc Information System (GIS) GRASS 4.0
software to measure the 10lal length of river occupied
frolIl lhe estuary to the furthest upstream record for
C<lch yeur. Therefore. if in year I cmbs were found
40 kill upSlream frum the e~lllary ill a pllnicular river
anti in year 2 were 60 km upstream from lhe estuary III the same river, the aver:tge anmla1 distance
of upstream migration is 30 km/year. 'nlis unit of
measuremelll was l:hoscn sim;e each populatioll found
furthcr up~trcall1 lhan any previously observed ones
must originale from the estuary dllc to its migration
betulViolif.
Size distribution data of the mitten crab populations ovcr sevcral years were l:ollectcd <lnd a wide
area of Gcrmany asscssed during the 1930s' inv<lsion
(for population distributions see Peters. 1933: Panning, 1938a: for migration. see Peters, 1933; Panning.
1938b). This information frolllthe major invasions in
northern Europe has becn anlllysed to describe the Iifehistory charaClcristics of lhis species. DUla from the
Lower Elbe from 1932-6 were pooled for each month
of the year to obtain average monthly size-class distributions of £ sinellsis. 11le dala (11=8736) were derived
from various sampling methods: slllali specimens (as
Migralory behaviour was assessed using calch (in kg)
from cmbs collecled in various Iypes of traps placed
at weirs to catch minen crabs during migrntion. These
were installed in the 1930s to reduce the mitten crJ.b
population wh.ich was a major pc::>l by lhis lime. Based
on the struclure of these tmp'>, the only crabs caught
\~ere those migrating across or around lhe weir (Panning. I 938b). Panning (1938b) used carapace marking
experiments to determine the downslream migratOr)'
speed of £'sillCn.\·is over variuu~ di"tunces. TIle recatch occurred eithcr in the crah tmps around a weir
or by fishermen.
Results
Sprem! m:ros.I" Con/inell/a! E!/mpe (//l(J StJlllhem
FrOJlCC
The first record of £. .l"iflensL~ was in 1912 in a tributary
of the Rivcr Weser in North Gennany (peters, 1933);
it was also found in the River Blbc from 1914 onwards
(both ri...·ers mn into the Nonh Sea only 60 kIn apart).
From thcre, il reached the Baltic Sea via the Kid Canal
in 1927 (Pctcrs, 1938a). Despite the fact that the number ofmiucn crabs in the Ballic remaincd much lower
than in Ihe river syslems along fhe North Sea, the species spread as far as Vyborg (Russia) and Finland by
1933. In the Netherlands. the first crab was recorded
in 1931 and the population had spread into mosl rivers
by 1936 (Kamps, 1937). It fCached Belgium in 1933
24
IlXXI
(Peters. 1938a and citations within: Lcloup, 1937).
France was invaded by 1930 (Hoestlandt. 1945) and
Denmark by 1927 (Rasmussen, 1987).
An important finding was also the extent of spread
upstream in somc rivers. E. sil/ensis had migrated
along the tributaries of the Elbe as far as Prague in
the Czech Republic (700 1.::111 inland) by 1932 (Peters.
19383). In the same year, it had reached 5 12 km along
the Rhine and. by 1934. it had spread 464 km upstream
in the Oder to Breslau. Analysis revealed that a separate invasion occurred along the river Gironde (19541960) in SOllthern Fr.mce. Previously, E. sinensis had
spread only as far as Le Havre (1943) along the French
Channel coast (Hoestlandt. 1959: Vincent, 1996). In
1954. specimens were caught in Nantes in the River
Loire estuary and the same year nellf Bordeaux in
Ihe estuary of the Gironde. Migrating upstream in
the Gironde. the mitten crab reached the artificial lagoons along the Mediterranean coast via canals by
1959, a distance of 504 Km (Petit & Mizoule, 1974).
These measurement, gave an average total di&tllllce
of upstream migration of 48 kmlyear between 19 I21927, 562 km/year for 1928-1939 and 69 km/year
in 1940-1955 for Northern Europe, and 104 kill/year
for Southern France (1954--1960) (Fig. 2). The distance recorded upstream for the nonhern European
and southern French river systems in each year of the
in\'ll,ion is shown in Figure 2. In Northern Europe
the Chinese mitten crab spread slowly o."cr a 15year period (1912-1927) but then &pread mpidly from
192810 1939. slowing aflerwards. Pan of the apparent slowing in the spread of mitten crabs in Northern
Europe after 1939 might be related to the reduced
monitoring effort during the war. In Southern France,
records show no establishment phase but a rather sharp
increase in spread between 1954 and 1960.
Obsen'atiolls 011 population distribution aJl(1
migrator)' lffitlem
The size-class distribution (Fig. 3) of mitten crabs
(both male and female) in the lower Elbe shows
the presence of at least two differelll cohorts during
spring: a larger/adult class (54.6 mOl ± 5.3 mm d for
January) disappearing in May. and a smaller/juvenile
class (29.8 mm ± 7.6 mm cI for January) decreasing
in size considerably by May (12.4 mm ± 3.4 lllm d).
The smaller cohort remained over the whole year. flucmating in mean size between 8.6 mm ± 3.5 mm cl
(September) and 20.7 mill ± 4.4 mmd (December). In
June, the larger cohort was found again in the estuary
10\100
-.+-,.,,-.,~
~
!
BOO
8000
1'000 §'
8000~!
S c 700
~!Wl
1~ soo
&IOOi./l
H~
-.
~H5
,
30001
.;:"'JCJ:l
6 200
20CC
1~ 1..,""""!!!.=11110
1>"9
1930
V
-_j_~1;~
1940
lnt"'l'O'l
,
1950
Figure 2 TIloI' cumulallV<: tOlal le<lgth of rh'ef (in k.m) OCCUpIed
hy Eriocheir ~i"e,ui~ from the esluary 10 lhe furthesl upSlream~.
cord for each year. melliourod using <A'Qgmphic tnfoml3tiOli SySlcm
(GIS) GRASS 4.0 softwaI\': The spread in Northern EtJrope ( _)
is plotted scparmely from the spread m Southern Fr:lJ1ce (t), NOll'
tile difftrelll <cates of distance of Ii"er in"alkd used for the lWO
im·aSlOIlS.
,,
•<
"
M
f f
M
r '"
[
M
,,i• " f
"
0
0
Tifftf
tf f T
ff
t
-
f
f f Tf
, , , , , , • ,
10
11
12
Figure J The size-<Iass distribution (± I SO) of Errochrir $;nrnsi~
the lower Elbc: per month as dclermim:d using modal progre.ssion
analysis (Bhan"eharya's IlIdhod) In tile FiSAT stock as~ssment
p;1oCbgc.
In
with a smaller carapace length (31.5 lrun ± 6.7 mm),
increasing gradually until December to 51.7 Illlll ± 4.9
mmcl.
TIle mean carap.1ce length of upstream-migrating
E. sinensis, measured using animals caught in traps
at weirs along the Weser and Elbe, showed an average
carapace length increase of 3 nun per 100 km upstream
from the estuary (Fig, 4) (Panning, [938b). This can be
used to estimate the annual growlh rate as 6.6 mm over
the annual migration distance of225 km (according to
Panning, 1938b). The annual growth rate estimated by
(Panning, 1938a) suggeSls an increase of about 12 mm
per year up to year4. Despite the discrepancy of Ihese
growlh rates. they are the only available estimates for
E. sinensis in Europe. The peak downstream migration
period at three locations (estuarine. 310 km upstream.
....
=
25
50
y • O.0295>c • 23 1<15
Discussion
f
150
200
250
300
kmfromltlll~
F'g"" <I 1lIe mean carapace length (in mm) of F.n·oc~lr sinmsls
IfUgnolUll upsunm along the RI,ws EJbe IIIld "~lGenn..nY'JOO
250
•
§
200
,~
150
100
50
0
I
1933
,
1935
1934
alion for two ri"ers out of many in Europe this species
has invaded. the lotal popul3tion must h3ve been "ery
large.
1936
V."
Fig..n 5. The annual eatch of Erioch~/r JIMlUiJ (in "'Nnc tons)
frOI11 tile RlI'cr Weser alld Elbe (Germ.any) during the in'OlSion of
Northern LUrope (1933-1936).
410 km upstream) was August-October. Absence of
a difference in migration period between these sites
can be explained using downstream marking experiments. These showed thaI mature animals can cover
large distances (400 km or more) within this 3-monlh
migration period, since analysis revealed an average
downstream migration sPCI..-<! of 11.5 kmlday (SD±
3.54) (Panning. 1938b).
Using the catch data from the rivers Elbe and
Wescr, population numbers can be estimated for the
mid 1930s; the peak catch occurred in 1936 with an
annual catch of mitten crnbs t()(alling 242 IOnnes in
Gennany alone (Panning. 1939b) (Fig. 5). Based on
an a'-ernge weight of 55 g per indIvidual from collec~
tions in the river TIuunes by Herberg (unpub!. dala),
total catch can be eslimalcd to be the equivalent of
4.4 million crabs. Considering this IS only an estim-
lhe most obvious fe.1ture of these resulls is the similar paltern in expansion in length of river colonised in
both Southern France and 'orthern Europe. In bolh
m-er syslems there is e\ idence of a sigmoidal relaTionship between length of nver iO\'aded and tllne.
Whilst the pattern may have been similar. comparison
of lhe rales of spread bclINCCll Northern Europe and
Southern frarx.:c indicates a much faster mtc of spread
during the eJl:ponentlal phase of the spread In 'orthern
Europe. These differences probably reflect differences
in the atructure of the 1"'0 ri\'er systems. In orthem
Europe. nvcr systems arc el(lensive. especially along
the North Sea coaST where the Rhine. Weser and Elbe
give access TO an extended nelwon. of tributaries as
well as to canals linking to other ri"ers. The outbreak
in Southern France was limited by the nature of the
Glconde systcm. which is less extenSive. Towards The
south. the Pyrenees fonn a n:llural barrier and, 10 the
north of the Gironde. relatively few large rivers run
into the Bay of Biscay.
1llc presence of a 15-)c:lr period of slow spread in
Germany :11 the e.'lrly slage of in\'asion can be imerpreted as an 'establishment' phase which is observed
commonly during invasions (Drake & Williamson.
1986). Absence of:l establishment phase in Southern
Fr.mcc mllY be due to a lack of data. or perhaps thaI the
Chinese millen crab population. at Ihis time already
present for over 40 years in Europe. was well acclimalised to European environmental conditions. Another
factor associated with a sharp population increase of
£. sinensis is periods of low rivcr flow as observed for
the U.K. in 1989-1992 (Attrill & lhomas, 1996).
The pattern of spread of the Chinese mitten erab
in Europe compares well with Ihe spread of the
European shore crab (Cmrinus maenas. L.) along the
Californian coost. 80th species have a similar pelagic larval period in their life cycles (ca. 60 days
in 20°C) (Anger. 1991; Cohen et aI., 1995). Obscn'atlons of the inva.<:ion rates of C. mOt:nas also
showed fluctuations (between 16 and 63 kmlyear) in
the rate of spread along tbe coast in different invasive
e\'ent5 (Grosholz & Ruiz. 1996 and citations within).
Grosholz. (1996). analysing the rate of spread for JO
invasions of the marine em'ironment, reported an aver-
•
26
age flltc of spread for marine il1Vadcrs of 50.7 krn/year
with individual values between 12 and 115 kill/year.
Estimates of the ratc of spread for three invertebrate
species (Balwills improl,jsus, Darwin.PotamoPY"8l1s
anripodarillnl, J.E Gray, Marenzelleria viridis. Verrill)
in the Baltic Sea range between 30 and 480 kmlycar
(Lcppaekoski & Olen in, 2000). The sigmoidal Tale of
spread has 'llsa been described in other invasions: the
spread of red decr (Cerous e!aplllls, L.) on the South
Island of New Zealand described a 40-year establishment phase before the area of invasion increased
rapidly, fJaucning off again afler ca. 20 years (Clarke.
1971).
Whilst the spread of E. sincllsis in Northern Europe
seems 10 originate from the Weser area. the question
arises as to the origin of the Chinese mitten crab at
this site. An introduction via ball:lst water from China
is the most likely vector. since there has been frequent
ship traffic from Hamburg (River Elbe) and Bremen
(River Wescr) to China since the end of the 19th century. The typical steamer on this routc had 15 ballast
water tanks, each holding 90 tonnes. with an inlakc
grid of 15 x 460 mm, large enough not only for larvac
but also juvenile mitten crabs. Peters (1933) mentions
the case of two adult millen crabs (40-50 llllll) which
were found in the sea chest of one of these vessels during its salvage. Introduction of the Chinese mitten crab
is considered as one of the very first known cases of a
s(X.'Cics being transponed via ballast water (Carlton.
1985).
It is likely that invasion of Southern France arose
in a similar way. The arrival of millen crabs in Southern France demonstrated a significant jump in spread
along the coast. Before 1954, the year the Chinese mitten cmb appeared in the Gironde and the Loire, it had
only been reponed from the Seine, a distance of 855
and 205 km along the coast, respectively. Considering
this distance, as well as the prevailing surface transport going northwards (Ellicn et aI., 2000; Alvarez et
aI., 2001), larval transport along the coast is highly
unlikely. The most likely vector is human-aided Transport, most likely via ship ballast water or in association
with the intensive oyster aquaculture along this coast.
Furthennore. the long time gap between the invasion
of the nonhern Atlamic coast of France. and that of The
Bay of Biscay. also suggests that a chance ship transpon across an expansion barrier was the mechanism of
introduction rathcr than natural spread.
The size·frequency distribution of crabs in the Elbe
eslUary over the year shows Ihe recruitment. departure
and anival of different cohorts of animals accord-
ing 10 Their life cycle. At the beginning of the year.
a 1arg~ and small cohort was observcd. TI1C Illrgcr
cohort comprised animals in excess of 43 mm. The
minimum size for sexual maturity of both sexes, and
arc 1Il0st likely animals frOI1l the previous autullln's
mating season which are either ovigerous female~ or
part of The small fraclion which survives the maTing
season (Peters. 1933). The size reduction of tlle juvcnile cohort from March to April coincided with the
pe:lk upstre:lllll1ligration period as determined by migration rates (data not shown). The carapace length
of juveniles leaving the estuary may indicate that the
smaller cohon observed in the Elbe (January-March)
consisted of year-elass I and 2. whereas from May
onwards. only smaller crabs of the year I cohon re·
mained ill the estuary and the year 2 cohort started
their migrtltion further up~trea/1l.
Fluctuation of the mean si",.e of the jU\'enilc cohan during the summer and autumn months could
be caused by twO separate factors. Firstly. the settlement of megalopa larvae from the water column
in July-Seplember (Schubert, 1938) at a lengTh of
5mm (Schnakenbeck, 1933) and, sccondly, continuo
ous growth of the previous year class. The larger size
class appearing in the Elbc as early as July may be
explained as being either animals which relllllined ill
the lower part of the estuary, or juveniles migrating
downstream to The estuary to become sexually mature.
Further increase in this cohort is caused by adult crabs
arriving downstream during the peak migratory period
(August-October). Analysis of catch numbers along
the Elbe and Weser showed that August to October
is the pe<lk downstream migr:lIion period in the River
Elbe (data not shown).
Recent invasions of Chinese millen cmbs in Great
Britain (Attrill & Thomas, 1996; Clark et al., 1998;
Herborg el al.. 2002) and San Francisco Bay (Rudnick
et aI., 2003) show similar patterns of spread to those
analysed here. There is also evidence that the invasion
process is ongoing, as recent findings of E. sinensis
in the Black Sea (Gomui et aI., 2002) and the Se.1 of
Azov (Murina & Antonovsky. 2(01) underline. The
study presented highlights the polential of major invasions of Esi/lensis. There has bcelllitlie description in
the historic work of the ecological impaci of invasions
such as those discussed here. Nevenheless. considering the size of the population at its peak, ecological
impacts may well have been considerable. What is required now is some quantification of these impacts, in
order that threats to biodiversity can be assessed and
27
requirements for management of the 111\ aSlOn problem
identified.
LMH would like to than},: the Faculty of Science. Agriculture and Engioc"Cring. Unhersity of Newcastle
(Swales Stutlcnl5hip) and the Esrnee f:llfbaim Found~
3110n (RG 01-1816) for their fin3nc131 suppon.
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Acknowlt.'dgcmenLs
dum:! speCIes in 1Cm:,trial and marine SYSlCrrIS. Erolog)'
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tbc: bmJpeaa grttll crab Corcinus MiMrtlU. BIOI. Con$cr\'. 78:
n:
922lloeglandl. II . 1~5. u crabe dtinois (E~lr .JlJInUU Mil Ed_)
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Hoesthmtll. H. 1959 R~lion l\Il:Iue:lk du crabe ChlDOIS
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1'i,lcul 19.1 S-13 (In French).
H)man~l. 1... J. wooS & T. Sas:tll. 1999 u"SOO$ rrom the home
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Ingle, R W, 1986. The Chmese nulten ~rnb I:.mxltelr s;nl"l$!s 3
continuous immigrant Lond. NllI. 65: 101-lOj
Jin. G, 7_ Li, & P. Xie, 2001. TIle 1:I<)....'lh pallen'S of jU\'elUle and
pm:ocioos ChmesC' millen lnb~. Eroc/tt!ir s;'W:n.lU (dl:Capoda.
grnpsid.1l. slocked m freshw:aler ~\..es or Chma CruSlaccalla 74
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Kamj);, L l-. 1937 De O1inee!iChe .... olhand mllm Nederland_
Akad. Prod Grooinllen: 1-112 (in DulCh)
Koba)"llSIu. S. ZOOI Nl:UOOIl) of !he J:tpanc:se mmen crab
EriocMir ~iro (de Hun) Benth R~ ~, 1-7,
~)"ashI. S .t: S MIU'UW'a, 1999. RqIRldueu\~ erolog} of tbc:
Japanese I1\IlIeIl <nil EriocIteir JiIPOtU'NI Ide Hunl • R:\''''''
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Lcloup, E.. 1937, Cootribulions 1l'Cludede la faunc beige VII _ La
ptOplIgllllOrl du cnbe chinois en 1k11!:lQue: pcndanl !'alIn&: 1936Bull \ill5.1I1$l. NlIlII' Belg. 13: 1-7 (in rrencJl)
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(Deupod;t Grarsidael reared in the lahnr.lIory H..l!01aender
Mtt'R'SUnll:n, jQ: 22J--252.
Murin;a, V V &: A G AnlOOO\·sl:)'. ZOOI ChIDl'!le crab. Enor~1r
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PeIen. N.• 1933. LcbcuskuDdlicba Ta1. In ~terS. N.• A. Panning
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AnL (l..c,pugl 59-155 (10 Ger-noanl
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Wollhandl:r3bbe (EnCJdterr S~IUU H.M -Ed.. ) UI E~ lID.
Jahre 1933 bu 1935. MIlL Hamh Zoot. MIlS. InsL 47: 1-31 (in
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Pc:Ien. R. 1938b. Neue: UDtenuehwlgn> I1ba die EnIIaiten der
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German).
28
N.. 193&. £ur
j'oopflunLullt\sblologlC dcr WollllamJkntlJlx.>
SctJubert. K.. 1938 Hiiutullll:. W""hSllIln IIml Aller ocr Woll·
(EriocMir ~"'CI>n.. II.M-l3dw). MllL Ilamb. Zoo! Mus. InsL 47:
112-128 (in Gcnnan).
I'ell•• G, & R r.hwule. 1974 F.n douZC;\l1S Ie 'crabe Chinois' n'a
pu riUSSIf son implanuuon dil/ls ~ lagulk5 du Languedoc. Vie
halld1:rabbt. Mill. lIamb. Zoo! Mus. InS! 47 83-104 (in
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Veldhuizen. T C &. S Sl.3ni<h, 1999 O'~I'\iev. of lhe life hiSlOl')'.
dl.mbullou. abu.nd.anl'c. and rmpllC'l of II~ Chmesc millen crab.
EnocMir S'~l.$. Oalir,..,ia {)q),)ltmenl of Waler Resources.
Emuonmemal ~J"\'Il'e5 OfI1ce.
ViOCWI, T.. 1996. Le crnbe CllillOis F",oclu'ir S"'''''n> H MIl"e·
l'ell'l'1i.
MJiJetJ
Sef C BIOI Terr. 2.3 181 186 {m rrmc:h}.
Rasmu~n. E..
('1I1;S'
1987. SlalUs O\'Cr uldh1ndsknbbens (Eriodwir sin-
udlJre.Jel'ie og f(Jl'l'~OIl~ , Damnark. Flora og Fauna 93
51-58 (in Danish).
ROOllld.. D, A, K Hrrb, K. F. Goml""'.t V H R<5h,2003 PatIcm.\ ;md pItlCC:Md of biolo&JaI II1\~JOn the C'bIllCSC mmcn
crab In S., FranclloCO 6a) BISIC" arvl I:rol .1 lin press).
Edwlll'ds. 18SJ (Cn15laca, Bnchrura) en xlne-Mariumc.
Fnoce, Ann IDSl~ 72 ISS 171 lin Freoch)