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1

1 An Ecopath with Ecosim model for the Pacific coast of eastern Japan:

2 describing the marine environment and its fisheries prior to the Great East

3 Japan earthquake

5 Shawn Booth1, William Walters2, Jeroen Steenbeek3, Villy Christensen4 and Sabine Charmasson5

6
7
8 1 Institut de Radioprotection et de Sûreté Nucléaire (IRSN) BP 3, 13115 St-Paul-les-Durance, France; shawnrgbooth@gmail.com

9 2 Penn State University, Department of Nuclear Engineering, 232 Hallowell Building; wjw@psu.edu

10 3 Ecopath International Initiative, Barcelona, Spain; jeroen.steenbeek@gmail.com

11 4 University of British Columbia, Institute for Oceans and Fisheries; v.christensen@oceans.ubc.ca

12 5 Institut de Radioprotection et de Sûreté Nucléaire (IRSN) BP 3, 13115 St-Paul-les-Durance, France; Sabine.Charmasson@ifremer:fr

13

14

15 ABSTRACT

16

17 A mass balance model of the marine ecosystems is presented for the four prefectures of

18 Japan prior to the Great East Japan earthquake. The Ecopath with Ecosim food web

19 modelling approach was used and its Ecotracer routine were used to simulate the

20 concentrations of 137Cs in the ocean and the biota in order to serve as baseline, to serve as a

21 marker for normal background conditions, and to estimate the initial conditions for impact

22 scenarios. Results of the Ecopath model were checked against pre-balance [PREBAL]

23 diagnostics for integrity. 137Cs was shown to bioaccumulate with increases in trophic level,

24 and has a tendency to biomagnify. Generally, for fish species, the dominant pathway of

25 contaminant accumulation was through diet, whereas for invertebrates it was through

26 respiration. Ecotracer was able to accurately model the expected concentrations in

27 organisms that had existing concentration ratio data from field measurements, and also

© 2020 published by Elsevier. This manuscript is made available under the Elsevier user license
https://www.elsevier.com/open-access/userlicense/1.0/
 2

28 make predictions on biota lacking concentration data. In 2010, using national fishery landing

29 statistics for the four prefectures and predicted whole body concentrations of 137Cs, it is

30 estimated that the apparent direct and indirect human food supply of 137Cs from fisheries to

31 the market was 46.1 MBq.

32

33

34 KEYWORDS: Marine ecosystem modelling, Ecopath with Ecosim, Ecotracer, Fukushima,

35 137Cesium

36

37

38 1. INTRODUCTION

39

40 The Great East Earthquake, the subsequent tsunami, and nuclear power accident in Japan on

41 11 March 2011 caused a significant loss of life, and also eroded a way of life for the people

42 who inhabited the coastal villages that were involved in fishing activities (Komatsu 2017).

43 The earthquake and tsunami caused significant damage to fishing infrastructure, vessels,

44 gear and fishing grounds (Japan Fishery Agency 2012) while the nuclear power accident

45 caused the release of large amounts of 137Cs into the ocean affecting the marine life after the

46 tsunami caused meltdowns at the Dai-ichi nuclear plant in Fukushima (Aoyama et al. 2016;

47 Kaeriyama 2017; Wada et al. 2016). Although other radionuclides were released in the ocean

48 such as 131I and 134Cs, it is 137Cs that is of concern because of its relatively long half-life of

49 around 30 years (International Atomic Energy Agency 2007), and the potential radiation

50 doses and associated risk to marine biota and humans from the consumption of seafood

51 (Fisher et al. 2013). In order to understand the direct ecological effects, it is necessary to
 3

52 examine the state of the marine environment prior to the accident and the role fishing

53 activities had on the marine environment before 2011. Presented here is an Ecopath with

54 Ecosim (EwE) ecosystem model for the marine waters of the four prefectures of Iwate,

55 Miyagi, Fukushima and Ibaraki that were most affected by the tsunami and subsequent

56 nuclear accident.

57

58 Our study area of approximately 220,000 km-2 focuses on the marine ecosystem for the year

59 2010 from the shoreline to the 200 nautical mile boundary of the exclusive economic zone

60 (EEZ), the region where both the coastal and offshore fisheries occur, and which is under the

61 influence of both the Kuroshio and Oyashio Currents. These two currents affect the

62 ecosystem and the fisheries in the area. When the cooler Oyashio Current expands

63 southward into the area there is an increase in biomass of, most notably sardines, and

64 Alaska Pollock. When the warmer waters of the Kuroshio Current dominate, there is an

65 increase in zooplankton biomass and a shift towards higher biomasses of Japanese common

66 squid and anchovy (Yatsu et al. 2013). The southward extension of the Oyashio Current

67 would also result in waters of a lower 137Cs concentration being brought into the area,

68 whereas the northern movement of the Kuroshio would bring in water with higher

69 concentrations of 137Cs into the region (Hirose and Aoyama 2003).

70

71 The modelled area generally has a narrow coastal shelf extending out to 40 km although

72 around Sendai Bay it extends to 70 km. The continental slope is also narrow and depths at

73 the Japanese trench can reach 9,000 m, but shallows to 5,000 m at the extent of the

74 exclusive economic zone where there is an abyssal plain (International Atomic Energy

75 Agency; IAEA 2015). However, in our study we limit the zone to an average of 1,000 m to
 4

76 reflect the taxonomic composition of sampled species that form the basis of the functional

77 groups in the Ecopath model.

78

79 Coastal fisheries occur within five kilometers from the shore, and offshore fisheries occur

80 from 5 kilometers of the shore to the boundary of the EEZ (Yamamoto 1985). Fisheries

81 landings from Japan’s Ministry of Agriculture, Forestry and Fisheries (MAFF 2014), and

82 monitoring data of cesium concentrations data in the ocean water (Nozaki et al. 1998; Hirose

83 and Aoyama 2003; Povinec et al. 2004) and in marine organisms (Kasamatsu and Ishikawa

84 1997; Yamada et al. 1999) were available for the study area. Although the earthquake

85 caused the release of large amounts of cesium into the ocean from direct and atmospheric

86 releases, there had been previous impacts from 137Cs on the marine environment from

87 nuclear testing during the 1960s and to a lesser extent from the Chernobyl event in 1986

88 (Duran et al. 2004; Smith et al. 2017).

89

90 In order to describe the state of the marine environment and the amounts of 137Cs present

91 prior to its release from the Fukushima Dai-ichi nuclear power plant, the EwE modelling

92 software was used. EwE has largely been used for describing marine ecosystems (e.g.,

93 Polovina 1984), assessing the state of marine ecosystems in comparison to earlier states

94 (e.g., Cox et al. 2002), providing information on emergent properties (e.g., Libralato et al.

95 2006), and being used as a fishery management tool to look at effects of changing

96 biomasses, linked to policy changes or climate change (Niiranen et al. 2013). However, here

97 we also deployed the Ecotracer contaminant module to describe the flow of 137Cs in the

98 organisms and environment. Ecotracer has been used to trace the flow of several pollutants
 5

99 in the marine environment including mercury (Booth and Zeller 2005), radionuclides in the

100 Baltic Sea (Sandberg et al. 2007), and in the Irish Sea (Tierney et al. 2017).

101

102 The aim of our study was to have a description of the biomass flows including the biomasses

103 at each trophic level, and the marine trophic index (MTI; Pauly and Watson 2005) to be able

104 to reflect the state of the marine ecosystem prior to the closure of the fishery after the

105 accident. To serve as a baseline prior to the nuclear accident, the concentration of 137Cs in

106 the environment and biota are estimated. As well, we examine the bioaccumulation of 137Cs

107 in the marine ecosystem and estimate the apparent human food supply of 137Cs to the

108 market in 2010 from fishery landings.

109

110 Despite recent corrections to the underlying computer code to the Ecotracer routine

111 (Walters and Christensen 2018), several suggestions are also made to improve the routine to

112 enhance its capabilities.

113

114

115 2. METHODS

116

117 We describe the marine food web through an Ecopath with Ecosim food web model

118 consisting of 55 biological groups and one detritus group. Included are the removal of 24

119 aquatic organisms reported in the 2010 capture fishery statistics (MAFF 2014). Functional

120 groups were considered to be coastal (e.g., shallow flounders), offshore (e.g.,

121 bathydemersals) or occurring throughout the entire area (e.g., mackerels). The Ecopath
 6

122 model was validated using pre-balance (PREBAL) diagnostics (Link 2010), and was

123 constructed adhering to best practices guidelines (Heymans et al. 2016).

124

125 We estimated starting 137Cs concentrations for water and biota in 2010 using data from the

126 1990s.. Water column concentrations, which resulted from nuclear bomb testing in the

127 1960s and to a lesser extent from the Chernobyl accident in 1986 (Smith et al. 2017), were

128 decay corrected to 2010. Additional marine inputs from low level radioactive effluents from

129 the nuclear plants in Fukushima are ignored as they were assumed to be negligible (Duran et

130 al. 2004). Similarly, scavenging of 137Cs by particulate matter was ignored because it is

131 considered to be limited to less than 1 percent in the open ocean (Aoyama and Hirose 1995),

132 and even after the Fukushima accident sediment trap data revealed that the removal rate of

133 137Cs on sinking particles was 1 to 2 percent per year (Buesseler et al. 2017).

134

135 For organisms, reported concentrations ratios from the 1990s were used, along with the

136 decay corrected water concentration to estimate concentrations in biota. Physical decay

137 rates based on the half-life of 137Cs were applied to both the water and functional groups.

138 The model was run forward in time using the Ecosim component to estimate potential

139 concentrations in functional groups and species that lacked such data under environmental

140 steady state conditions. Given that 137Cs concentrations are considered to be well mixed and

141 spatially homogenous, we did not undertake a spatial-temporal assessment.

142

143 It is outside the scope of this work to describe in detail all the specific numerical methods

144 contained within the EwE modeling framework. However, a brief description of the master
 7

145 equation used in the Ecopath component is given, as well as for Ecotracer, the two main

146 components used within this study.

147

148 2.1 Ecopath

149

150 Ecopath with Ecosim uses a trophic mass balance approach for the Ecopath component,

151 which in its simplest formulation, ensures that the amount of consumption and the fisheries

152 yield on a group/species does not exceed its production (Polovina 1984). To account for

153 biomass changes, such as those from regime shifts, a biomass accumulation term can be

154 used; as well, if necessary, immigration and emigration can be dealt with using net

155 migration. Consumption rates of predators are used along with diet compositions of prey

156 items to estimate the amount of consumption of prey items, and mortality rates from non-

157 predation events (such as old age and disease) are incorporated as other mortality

158 (Christensen and Walters 2004). Thus, the production of a group/species (Pi) can be defined

159 as,

160 Equation 1)

161 = + 2 + + +

162

163 where Yi is the fisheries catch rate (t·km-2·year-1) of i, Bi is the biomass (t·km-2) for group i,

164 M2i is the total predation rate (year-1) for group i, Ei represents the net migration rate

165 (emigration – immigration; t·km-2·year-1), BAi is the biomass accumulation rate for group i,

166 MOi is the other mortality rate (year-1) for group i. Another form of the equation

167 (Christensen et al. 2005), which is useful in terms of the input requirements for the EwE
 8

168 modeling interface is to re-express Equation 1 in terms of the input parameters required for

169 EwE as ,

170 Equation 2)

171 ∗ ∗ = ∗ ∗ + + +

172

173 where (P/B)i is the production to biomass ratio for group i, EEi represents the ecotrophic

174 efficiency or the amount of production used within the system and is a value between 0 and

175 1, Bj is the biomass of each predator group feeding on group i, (Q/B)j is the consumption to

176 biomass ratio of each predator group feeding on group i, and DCji is the fraction of prey

177 group i in the diet of each predator group j. In this form, if the emigration and accumulation

178 terms are ignored, then it is conveniently seen that, in essence, the production of a group is

179 equal to the consumption (plus fisheries) of that group.

180

181 For each group in Ecopath, the diet composition and any fishery catches must be entered,

182 and, depending on the state of the ecosystem or type of organisms, estimates of biomass

183 accumulation and emigration can be entered. Also, all but one of the remaining parameters

184 (i.e., B, P/B, EE, and Q/B) must be entered for each group, as the mass balance routine can

185 estimate the missing term (Christensen et al. 2005).

186

187 The MTI reflecting the state of the fishery landings of high trophic level fishes in 2010 is

188 given for the cut-off point at TL 3.25 (3.25MTI) as suggested in Pauly and Watson (2005). The

189 MTI was calculated using the weighted mean catch by trophic level, i.e., MTI = ∑(TLi*Yi)/∑Yi

190 where TLi is the trophic level of a group and Yi is the reported landings.
 9

191

192 2.1.1 Groups

193

194 The species and groups represented in the model are formed from the types of information

195 available from a database created from the monitoring on radioactivity level in fisheries

196 products which covers the years 2011-2016 and provides information on 137Cs levels in

197 marine aquatic organisms after the Fukushima accident (Japan Fisheries Agency 2017). The

198 263 species sampled in the four prefectures were retained as individual groups or were

199 placed in functional groups for modeling purposes. Contaminant, dietary, landings, and

200 habitat data were used for this purpose.

201

202 We combined species of similar ecological roles into functional groups using a scoring

203 system based on the number of 137Cs samples taken per year. An index of 1 was given to

204 those species which were sampled more than twice in each year in a prefecture, and a score

205 of 4 was given to species not sampled in a prefecture. A score of 2 was given to those species

206 that were sampled once each year in each prefecture, and a 3 for being sampled at least

207 once over the sampling period. Species that scored a value between 1 and 2.25 were

208 considered to be candidates as individual species in the model after appraising fisheries

209 statistics, and the dietary data available.

210

211 Japanese fisheries statistics are mostly reported at group level (e.g., flatfishes); however,

212 several species relevant in the model area are reported individually including tunas (Thunnus

213 orientalis, T. alalunga, T. obesus, and T. albacares), skipjack (Katsuwonus pelamis), round

214 herring (Etrumeus teres), anchovy (Engraulis japonicus), saury (Cololabis saira), cod (Gadus
 10

215 macrocephalus), Alaska pollock (Gadus chalcogramma), Atka mackerel (Pleurogrammus

216 monopterygius), yellowtails (Seriola quinqeradiata) and the common scallop (Patinopectin

217 yessoensis). Of these independently reported species in the fishery statistics, only cod scored

218 an index of one. However, olive flounder, which is not reported independently in the

219 fisheries statistics, also scored an index of one, had good data concerning biomass (Shibata

220 et al. 2017), and thus was represented individually in the model. Landings reported at the

221 group level were disaggregated assuming the catches in the group were of high, medium or

222 low levels.

223

224 Other individual fish species in the model were marbled flounder (Pleuronectes yokohamae),

225 flathead flounder (Hippoglossoides dubius), Japanese jack mackerel (Trachurus japonicus),

226 and fat greenling (Hexagrammos otakii). These species had a score of 1.5, 2.0, 2.0, and 2.25

227 respectively. Marbled and flathead flounder were split from the group ‘Flatfishes’ as

228 reported in the Japanese fishery statistics and both had seasonal diet data. Japanese jack

229 mackerel is reported separately in the fisheries statistics, and diet data were available from

230 Japan (Tanaka et al. 2006). Fat greenling had both dietary data and a detailed study of 137Cs

231 present in the muscle tissues near the accident site (Shigenobu et al. 2014). Alaska pollock

232 was also chosen because of it being reported separately in official statistics, and it had a

233 detailed diet study by depth in Japan (Yamamura et al. 2002).

234

235 The remaining species data from the database were grouped to form functional groups.

236 Taxonomy, diet, depth of occurrence, and size all played a role in forming the groups.

237 Primary producers are represented by phytoplankton and algae, and zooplankton consist of

238 three types based on diet—herbivorous (copepods), omnivorous (mysids, ostracods and
 11

239 some non-gammarid amphipods), and carnivorous (chaetognaths, hyperiidea, and

240 coelenterates). A group of detritivores are also included that consist of isopods, gammarids,

241 cumaceans and cladocerans. The krill species, Euphausia pacifica, is allocated to its own

242 group and includes a fishery of approximately 5,600 tonnes.

243

244 A simple representation of the macrobenthic compartment comprises nine functional groups

245 and includes bivalves, gastropods, annelids, sea urchins, sea cucumbers, shrimp (omnivores

246 and predators), seastars, and the group crabs and lobsters.

247

248 2.1.2 Diet matrix

249

250 Several factors are important in the trophic modelling of marine organisms including

251 seasonal, spatial and foraging depth variations. When possible, we used local dietary data

252 that was collected seasonally and by depth as different prey items can be associated by both

253 factors. Seasonality can affect prey species availability and, for non-pelagic species, depth

254 can be an important factor in determining diet due to it affecting age stratification. For most

255 functional groups quantitative data concerning the wet weight were used, but for some

256 groups diets were only qualitatively described (Supplementary data 1). Unassimilated

257 consumption, the portion of a prey item not assimilated, was assumed to be 0.2 for all

258 groups, excluding herbivorous zooplankton and omnivorous zooplankton which were set to

259 0.4 and 0.3, respectively. This was done to reflect the lower food quality (i.e., phytoplankton)

260 in the diet.

261

262 2.1.3 Ecopath Parameters (B, P/B, Q/B and EE)

 12

263

264 Biomass estimates were made for 19 groups based on stock assessments used for a model of

265 the northwestern Pacific (e.g., Pacific cod; Watari et al. 2018) scaled to the local area or, for

266 zooplankton groups, on previously published estimates on production (Liu et al. 2005)

267 combined with species composition data (Ikeda et al. 2008). For groups without biomass

268 data, it was assumed that the ecotrophic efficiencies (EE) were 0.95 which enabled the mass

269 balance routine to estimate the biomass. The ecotrophic efficiency of a group describes the

270 amount of production used within the modeled ecosystem; for groups with B, P/B and Q/B

271 data the EE is estimated by the mass balancing routine.

272

273 The production-to-biomass ratio for fish groups can be considered to be the natural

274 mortality rate plus the fishing mortality rate (i.e., P/B = F + M; Allen 1971). Values for fish

275 groups were taken or calculated from a variety of sources including Watari et al. (2018),

276 from stock assessments, using estimates of natural and fishing mortality, or by assuming a

277 production-to-consumption ratio (P/Q) ranging between 0.1 and 0.3 which is consistent with

278 bioenergetics theory. In cases where the natural mortality was used in conjunction with

279 fishing mortality, the estimate of M was based on an empirical estimate (Pauly 1980). Values

280 for invertebrates were either taken from an Ecopath model for the Bering Sea (Aydin et al.

281 2002), assuming a P/Q value of 0.2, or from local Japanese data (Ikeda et al. 2008; Mori et al.

282 2009; Taki 2006). Phytoplankton values were based on Liu et al. (2004), and algae based on

283 the range reported by Mann (1973) and Schwinghamer et al. (1989). Detritus was based on

284 an empirical formula (Pauly et al. 1993) assuming a euphotic depth of 30 meters.

285
 13

286 The consumption-to-biomass ratio (Q/B) defines the amount of food consumed by a group.

287 Empirical estimates of Q/B for fish groups rely on using estimates of weight (W∞), the tail’s

288 aspect ratio and temperature (Heymans et al. 2016). The aspect ratios for tails were found

289 for the groups mackerels (Westneat and Wainwright 2001), medium pelagics (Collette 1978;

290 Westneat and Wainwright 2001) and small pelagics (Palomares and Pauly 1998; Westneat

291 and Wainwright 2001) and these were used to estimate the Q/B on the empirical

292 relationship. For the remaining fish groups, the life history tool and the average depth of

293 occurrence in Fishbase (Froese and Pauly 2018) were used along with a temperature profile

294 (Fujii and Kamachi 2003) to estimate Q/B. For zooplankton and krill, it was assumed that the

295 P/Q was between 0.2 and 0.3; estimates for shrimp were taken from Coll et al. (2007), and

296 the rest were taken from a model for the Bering Sea (Aydin et al. 2002).

297

298 The widespread use of ecological network models, such as EwE, of variable quality has led to

299 the call for a measure of quality control (Link 2010) in order to improve model parameters

300 and for aiding management bodies in the evaluation of model integrity. Several indicators of

301 the PREBAL diagnostics are used to check for model integrity (Heymans et al. 2016; Link

302 2010). Several of these diagnostics are presented including those for P/B, the gross food

303 conversion efficiency (P/Q), biomass and Q/B across trophic levels, and ratios of biomass. All

304 parameter estimates for B, P/B, and PREBAL diagnostics are found in the supplementary

305 material (Supplementary data 1).

306

307 2.2 Ecotracer

308
 14

309 The Ecotracer sub-routine can be used once an Ecopath model is in mass balance. Ecotracer

310 is adaptable to using different units depending on the contaminant and the concentrations

311 found in the natural environment and the type of data available. Thus, the sub-routine can

312 use different measurements including mass (e.g., grams) and radioactivity (Bq), but

313 considers the environment as a spatial area (i.e., km2) to match the underlying Ecopath

314 model inputs. An important consequence of this capability is the necessity to maintain

315 consistency in units in the environmental and biological compartments. In this model the

316 contaminant measure in the environment occurs in a spatial area (i.e., Bq·km-2) whereas the

317 radioactivity in biological groups are entered as amounts (Bq), and can be transformed into

318 group biomass concentrations (i.e, Bq·t-1) by the program.

319

320 Ecotracer tracks the flow of a contaminant in the environment and the biological groups

321 represented in a model through time (Christensen et al. 2005; Walters and Christensen

322 2018). The amount of a contaminant in a group, in this case 137Cs, is dependent on the

323 intakes and losses. Intake amounts come from direct uptake rates (i.e., respiration), the

324 fraction retained from trophic interactions, and immigration. Losses arise from natural decay

325 processes, metabolic losses and emigration.

326

327 Ecotracer is based on a standard dynamic transfer model (e.g., Landrum et al. 1992;

328 Thomann 1981) i.e.,

329 Equation 3)

)
330 = α − β! C )

331
 15

332 where αi represents the gains from direct uptake from the environment plus the gains from

333 diet, and βi represents the losses due to physical decay and elimination rates, which are

334 dependent upon the group’s concentration Ci.

335

336 However, Ecotracer expands upon the standard dynamic transfer model by including several

337 other terms important in an ecosystem context including migration, predatory losses, other

338 mortality, and fisheries. Another important difference is that for biological compartments,

339 the contaminant level is based on the amount of contaminant in a dietary item (i.e., Bq)

340 rather than the concentration (Bq·t-1), and the uptake rate from food is dependent upon the

341 parameters used in the Ecopath model to describe the amount taken in from all dietary

342 sources. Thus, the gains (Bq∙year-1) to a group are dependent upon a direct uptake rate, its

343 diet and immigration, i.e.,

344 Equation 4)

345 α =# $ + ∗ + %&
'()*

346

347 where, Co represents the environmental concentration (Bq·km-2), Bi is the biomass (t) of

348 group i, μi is the direct absorption rate proportionality co-efficient linking the direct uptake

349 rate (Bq·km-2·yr−1) to the parameters of biomass (t·km-2) and environmental concentration

350 (Bq·km-2), AEi is the assimilation efficiency and is the fraction of 137Cs within food items that

351 is assimilated, Qji is the consumption rate (t·year-1) of group j by group i, Aj is the amount

352 (Bq) of the contaminant present in group j and Bj is the biomass of prey group j (t), ci is the

353 group biomass concentration (Bq·t-1), and I is the immigrating biomass (t·year-1).

354
 16

355 The losses from a group (βi Ai) are attributed to predation, fisheries, other mortality,

356 excretion and decay, i.e.,

357 Equation 5)

358 , = +- + + +. + )
'()/

359

360 where Qij is the rate of consumption (t·year-1) of group i due to predation by j, Fi is the fishing

361 mortality rate (year-1), MOi (year-1) is the other mortality rate (i.e., non-predation mortality),

362 Ei is the emigrating biomass rate (year-1), mi (year-1) is the excretion and/or metabolic rate,

363 and di (year-1) is the physical decay rate. These rates are multiplied by Ai the amount of

364 contaminant (Bq) in each group i. Excretory products that are released from tissues to the

365 environment are added to the environmental concentration.

366

367 The Ecotracer approach is dynamic and extends the basic concentration ratio (CR) approach,

368 but the CR approach is contained within it as,

369 Equation 6)

1 2 # + 0
0 = =
3 + .
370
$

371

372 The environment is also dependent on gains and losses taking place in the environment.

373 Gains originate from the release of contaminants into the environment as a base inflow rate,

374 and from the excretion from organisms. Losses originate from the direct uptake by

375 organisms from the environment, physical decay rates, and base volume exchange. Thus, the

376 environmental compartment can be modeled as,

 17

377 Equation 7)

$ )
378 = 4 − , $ )

379

380 where αi represents the gains in the environment and βi represents the losses to the

381 environment. In Ecospace, the gains are from both additional inputs into the environment,

382 and from biota such that,

383 Equation 8)

384 4 = &+ .

385

386 where BI is the base inflow rate (Bq·km-2·year-1) to the environment, and miAi are the

387 excretory products from all organisms within the environment. Losses from the environment

388 are due to biological, physical and environmental volume changes, such that,

389 Equation 9)

390 , $ = +5) $ + # $

391

392 where di represents the decay rate (year-1), and Vi represents the base volume exchange loss

393 (year-1). Temporal changes through the environmental concentration Co can be made by

394 applying a forcing function to the base inflow rate or through a contaminant concentration

395 driver file.

396

397 The amount of contaminant in the detritus compartment originates from the unassimilated

398 consumption resulting from predation, as well as non-predation mortality. Thus, groups
 18

399 feeding on detritus will have exposure levels associated with the contributions from the

400 fraction of unassimilated consumption from all groups. Initial concentrations in the biota and

401 environment are also input parameters to be used if data is available.

402

403 2.2.1 Ecotracer Input

404

405 The initial environmental concentration was based on a dataset covering the North Pacific

406 Ocean. The 137Cs surface water concentration of 2.1 Bq∙m-3 for 1998, which represents the

407 mid-latitude areas of the western Pacific Ocean including the southern half of the Sea of

408 Japan, was used after it was decay corrected to the year 2010 (Box 2; Hirose and Aoyama

409 2003). A depth profile of 137Cs for the Japanes trench (Nozaki et al. 1998) formed the basis

410 for the starting concentrations. The depth profile curve was decay corrected to the 2010

411 surface water concentration value, and then an average value was calculated from 0 to 1000

412 m since below this depth 137Cs concentrations approached zero. .

413

414 For functional groups with reported CRs, initial amounts at the start of the simulation (i.e.,

415 Bq) were based on the CR, estimated water concentrations, and the estimated biomass of

416 the various functional groups. For fish groups, concentration ratios (CRs) for taxa in Japan

417 were used (Kasamatsu and Ishikawa 1997; Yamada et al. 1999) with the estimated water

418 concentration from surface waters to 246 m since the reported CRs were all for coastal

419 species and this depth is a typical water depth boundary between coastal waters and the

420 continental slope. For macroalgae, CR data was available for the study area (Tateda and

421 Koyanagi 1996) and for phytoplankton the IAEA weighted arithmetic mean was used (IAEA

422 2014). For herbivorous zooplankton, an average CR (8.0) was used from data collected in the
 19

423 western North Pacific Ocean from samples containing greater than 89 % of their biomass as

424 copepods (Kaeriyama et al. 2008). For omnivorous zooplankton the IAEA recommended

425 value of 40 was used (IAEA 2004), and for carnivorous zooplankton a CR was estimated so

426 that the average value for all zooplankton groups was the IAEA recommended value.

427

428 Functional groups without reported CRs had an initial amount of zero with the resulting

429 concentration being estimated by the Ecotracer routine. Groups in the model with and

430 without a reported CR were checked to see if they had a normal distribution using the

431 Shapiro-Wilk’s test and the two medians were compared using a Welch’s t-test using RStudio

432 (v 3.5.1). To evaluate Ecotracer, the observed activity values of groups with reported CR

433 values were plotted against the predicted values for the same groups and the root mean

434 square error is given (Piñeiro et al. 2008).

435

436 The direct absorption rate for the two primary producers were based on their CRi, Pi/Bi ratio,

437 excretion rate (mi), and the physical decay rate (di) of 137Cs assuming equilibrium conditions

438 (Walters and Christensen 2018) such that,

439 Equation 10)

440 # = 0 ∗ +. + )

441

442 where the P/B ratio for algae includes the harvest rate of seaweeds. This equation can be

443 derived by setting the derivative of Eq. 3 to zero, substituting the definition of CRi, and

444 solving for μi.

445
 20

446 For consumers, the direct absorption rate must also account for predation losses and under

447 equilibrium conditions it can be calculated as (Walters and Christensen 2018),

448 Equation 11)

∑
∗∑
'()*
67 ,)8 '()/ + +. + :−7 :;
# =
∗
449
$

450 where the first term in the numerator represents the losses due to predation (Qij/Bi), total

451 mortality (Pi/Bi), excretion (mi) and decay (di), and the second term represents the gains

452 from consumption of prey items.

453

454 The proportion of contaminant excreted (i.e., 1-AEi) is the amount of 137Cs that does not

455 enter tissues after ingestion. Fish species were assumed to have an AE of 0.8, which is

456 slightly higher than that corresponding to laboratory experiments undertaken with juvenile

457 fishes, but within the reported ranges (0.54-0.88) undertaken with juvenile fishes (Mathews

458 and Fisher 2008; Mathews et al. 2008; Pan and Wang 2016; Wang et al. 2016; Zhao et al.

459 2001), as data for adult fishes from laboratoty experiments are lacking. For 6 groups

460 (mackerels, piscivorous reef fish, pelagic neritic piscivores, squid, small and medium

461 pelagics), the direct absorption rate (i.e., μi) was set to zero, and equation 11 was solved for

462 the AE as the losses were less than the gains from dietary items.

463

464 For invertebrates, the reported AE of 0.16 were used for annelids (Wang et al. 2016) and

465 bivalves (Pan et al. 2016), and 0.5 for gastropods (Wang et al. 2000), crabs and lobsters

466 (Wang et al. 2016); it was assumed that sea urchins, sea stars, and octopus had an AE of 0.5.

467 Herbivorous and omnivorous zooplankton have an AE 0.65 (Mathews and Fisher 2008) and

468 omnivorous shrimp, krill, detritivores, and sea cucumbers were assumed to have the same
 21

469 value. For carnivorous zooplankton and predatory shrimp, an AE of 0.7 was used under the

470 assumption that their prey were of higher food quality. Primary producers were assigned a

471 value of one.

472

473 The physical decay rate is based on the radioactive half-life of 137Cs of 30.09 years (IAEA

474 2007). Excretion rates from a variety of lab studies were transformed from daily rates to

475 yearly rates and, for primary producers, an excretion rate from a model for the same area

476 was used (Belharet et al. 2016). The reported lab data was used to perform a regression

477 analysis using trophic level (TL) as the predictor variable to estimate the excretion rates for

478 all groups (Supplementary data 2).

479

480 2.3 Contaminant risk

481

482 For all groups in the model Ecotracer is able to estimate concentrations or burdens, leading

483 to the ability to estimate risk through time through such metrics as biomagnification factors,

484 and to also make comparisons to regulatory limits. Prior to the accident, the regulatory limit

485 for 137Cs in seafood products in Japan was 500 Bq∙kg-1, but was lowered to 100 Bq∙kg-1 one

486 year after the accident (Povinec et al. 2013). We estimate the apparent human food supply

487 of 137Cs, from fishery landings and the measured or estimated concentrations of 137Cs in the

488 group from fisheries landings (Yi) and the 137Cs concentration in group i (Csi,), as ∑Yi*Csi,.

489

490 In order to assess the behavior of 137Cs in the ecosystem, we examine its ability to

491 bioaccumulate using a biomagnification factor (BMF), i.e., a predator’s concentration of 137Cs

492 relative to its prey as defined by Mackay et al. (2018). For each group, we calculate the BMF
 22

493 as [Predi]/∑DCji*[Preyj], where [Predi] is the concentration of 137Cs in predator i, DCji is the

494 diet fraction of prey item j in the diet of predator i, and the concentration of 137Cs in each

495 prey item j [ Preyj].

496

497

498 3. RESULTS

499

500 3.1 Ecopath

501

502 The calculated biomass from the model representing the EEZ of all four prefectures in

503 eastern Japan (Iwate, Miyagi, Fukushima and Ibaraki) from the coast to the exclusive

504 economic zone comprising 55 biological groups plus detritus suggests that the combined

505 biomass was 391.7 t·km-2. Of the 55 biological groups large sharks and large pelagics

506 occupied the highest trophic level at 4.8 (Figure 1). . Biomass totals are 12.8 t·km-2 for

507 primary producers (macroalgae and phytoplankton), 181.8 t·km-2 for groups at the second

508 trophic level (2.00 to 2.86), while those at trophic level 3 (trophic levels 3.00 to 3.87) and 4

509 (trophic levels 4.00 to 4.78) have 48.3 t·km-2 and 2.8 t·km-2, respectively. Detritus has a mass

510 of 146.0 t·km-2. . The lower biomass of primary producers compared to the biomass at the

511 second trophic level is possible because of the much higher production rate of

512 phytoplankton (Table 1).

513

514 Total biomass production is estimated to be 3,160 t·km-2·year-1 with primary producers

515 accounting for 72.6 per cent. Groups in trophic level 2 account for 24.6 per cent of the yearly

516 production, those at TL 3 account for 2.7 per cent, while groups at TL 4 account for less than
 23

517 0.05 per cent of the production. The total capture fishery landings within the model area are

518 594,000 tonnes (2.70 t·km-2), and were dominated by three species with the largest reported

519 landings being mackerels (146,000 t), saury (64,000 t), and anchovy (54,000 t) which

520 together provide 50 per cent of the landings. The 3.25MTIfor fisheries landings of 496,508 t is

521 3.81.

522

523 The thermodynamic constraints (Heymans et al. 2016) for ecotrophic efficiency (EE < 1.0),

524 and gross food efficiencies (0.1 < P/Q < 0.3) are both met (Table 1). Net efficiencies (NE >

525 P/Q), respiration to assimilated biomass ratio (RA/RB < 1.0), respiration to biomass ratio

526 (RA/B), and the production to respiration ratio (P/RA < 1.0) are also within the range of

527 expected values (Supplementary data 1). The respiration to biomass ratio is expected to be

528 highly variable and generally they range from 1 – 100 year-1 depending upon the group, and

529 here, they fall between 0.89 for large sharks and 24.48 for krill. EEs for groups without

530 biomass estimates were set to 0.95, and for groups that had their EE estimated the values

531 ranged from 0.59 to 0.96 (Table 1).

532

533 PREBAL diagnostics (Supplementary data 1) indicate that the rate of decline in biomass from

534 the highest TL to the lowest TL declines at 9.8 percent, and the biomass amounts across all

535 TLs spans 6 orders of magnitude both of which are in agreement with PREBAL criteria (Link

536 2010). . However, in terms of the actual biomass levels for individual groups, there is

537 considerable deviation from the trend line: pelagic neritic piscivores, deep-water soles,

538 halibut and flathead flounder are well below the trend line, whereas groups such as

539 herbivorous and carnivorous zooplankton, squids, medium pelagics, and Pacific cod are

540 some of the groups that are above the trend line (Supplementary data 1). The P/B and the
 24

541 Q/B follow a decreasing trend from low to high trophic levels indicating that these vital rates

542 of the predators are generally lower than their prey. Deviations from the P/B and Q/B trend

543 line do not correspond to vital rates being higher than those of their prey, but rather involve

544 specific trophic interactions.

545

546 3.2 Ecotracer

547

548 In 2010, the concentration of 137Cs in surface waters to 246 m depth is estimated to be 1.5

549 Bq·m-3 while the average concentration from surface waters to the 1000 m depth is

550 estimated to be 0.5 Bq·m-3. Since EwE uses a 2 dimensional spatial representation, this

551 average amount was expressed on an area basis (i.e., 0.5 Bq·m-2), and scaled up to Bq·km-2.

552 Thus, the initial environmental concentrations was set to 504 MBq·km-2. To maintain

553 environmental equilibrium and in order to estimate the potential concentrations in biota in

554 2010, a base inflow rate of 11 MBq·km-2 was used to offset the physical decay rate of

555 0.023·year-1 (i.e., a half-life of 30.09 years). This spin-up to equilibrium was required in order

556 to allow the model to estimate 137Cs concentrations for all groups. Further, as reflected by

557 the P/B, groups in this model have a high turnover rate with no long-lived species, such as

558 whales being represented; thus, historical exposures to the small changes in water

559 concentrations due to physical decay process would have little effect on the resulting

560 concentrations in biota.

561

562 Direct absorption rates (μi) ranged between 6.38 x 10-5 and 1.27 x 10-2 km2·t-1·year-1. Lower

563 direct absorption rates were generally associated with high TL organisms and low TL

564 organisms had higher values (Table 2). For groups with direct absorption rates (μi) of zero,
 25

565 AEs were 10.2 % (squids) to 78.1 % (mackerels). It was chosen to alter the AEs rather than

566 the excretion rates (mi) as regression analysis of AEs presented in the literature did not show

567 a clear trend (results not shown), whereas a decreasing trend of excretion rates was found

568 with trophic level (Supplementary data 2).

569

570 The importance of the direct absorption rates can also be interpreted by the reflection of

571 their contribution to a group’s uptake of 137Cs from diet and respiration. Respiration

572 contributes on average 89.3 percent to the uptake of 137Cs for invertebrates, excluding the

573 groups octopus (19.8%), crabs and lobsters (46.9%), and squids (0 %). For fish groups, diet is

574 more important with 57.1 percent of the uptake of 137Cs originating from dietary sources

575 (range: 13.9 %– 100 %; Table 2).

576

577 Concentration ratios based on the surface water concentration of 1.5 Bq·m-3 for

578 phytoplankton and algae are 8.4, and 18.2, respectively. The average CR for successive TLs is

579 25.4 (range: 7.0-39.7) for TL 2, 46.6 (range: 8.8-94.3) for TL 3, and for TL 4 the average CR is

580 64.8 (range: 33.9-130.6). Groups with a reported CR had a median value of 44.9 when the

581 outlier is excluded, and those groups without CRs had a median value of 43.6 (Figure 2). Both

582 groups were normally distributed using the Shapiro-Wilk’s test (reported: w = 0.956, p =

583 0.299; unreported: w = 0.945, p = 0.133), and there was no difference between the means (p

584 = 0.55) when the outlier is included. For those groups with a reported CR, Ecotracer

585 estimated the observed versus predicted amounts (Bq) having a root mean squared

586 deviation (RMSD) of 23.09 (Figure 3) implying that Ecotracer has the ability to reproduce the

587 observed values well.

588
 26

589 Biomass concentrations range from 10.5 Bq·t-1 in seastars to 195.9 Bq·t-1 in the functional

590 group skates and rays, while detritus is estimated to have a concentration of 6.2 Bq·t-1. There

591 is a general increasing trend in the biomass concentrations of 137Cs with trophic level (Figure

592 4) with TL explaining 52 per cent of the variation of 137Cs concentrations (R2 = 0.5171, p <<

593 0.001). Functional groups of TL 1, including detritus, have an average concentration of 15.4

594 Bq·t-1, TL 2 have an average concentration of 38.0 Bq·t-1, TL 3 have an average concentration

595 of 69.9 Bq·t-1, and organisms at TL 4 have an average concentration of 97.2 Bq·t-1,

596 representing a 6-fold increase from TL 1 to TL 4.

597

598 3.3 Contaminant risk

599

600 In 2010, prior to the accident, all seafood products were well below the regulatory limit for

601 137Cs of 500 Bq∙kg-1 (Table 2). The apparent supply to the market destined for human

602 consumption on a whole wet weight basis of 137Cs is 46.1 MBq based on the reported

603 landings and concentrations of 137Cs estimated to be in the seafood products. 137Cs in the

604 ecosystem bioaccumulated and had a tendency to biomagnify, but not for all groups (Table

605 2). The average biomagnification factor for all groups was 1.92. However, 8 groups did not

606 have a biomagnification factor above 1, indicating that in these groups biomagnification did

607 not occur. The lowest values for invertebrate were for squid (0.18), crabs and lobsters (0.40),

608 seastars (0.47), and herbivorous zooplankton (0.99). The four fish groups were generally

609 closer to a value of 1 with forage fish feeding primarily on copepods have a value of 0.7, and

610 the remaining groups being above 0.90 (mackerels, omnivorous bathydemersals, and pelagic

611 neritic planktivores feeding on plankton).

612
 27

613

614 4. DISCUSSION

615

616 It was the intent here to model the marine ecosystem of eastern Japan surrounding the

617 Fukushima nuclear power plant prior to the nuclear accident to serve as a baseline. To do

618 this, an Ecopath model was constructed with the information that was publically available.

619 Although sparse data was available for the same year for the same species, it is encouraging

620 that most of the PREBAL diagnostics and indices of best practices fall within expected

621 values.. Data concerning biomass, diets and fishing effort for stocks within the area would be

622 valuable for improving the model and, if made available, it would be worthwhile to do a

623 model pedigree and time series fitting.

624

625

626 The purpose of the Ecopath model was to trace the concentration of 137Cs in the marine

627 ecosystem in the water and the organisms using the newly modified Ecotracer routine. The

628 model results showed that there was a general increase in the concentration of organisms

629 with increasing TL. However, in some cases a group at a higher trophic level does not have

630 higher concentrations than a group at a lower trophic level. For example, Alaska pollock (TL

631 3.75) has a predicted concentration of 141.5 Bq·t-1, whereas Pacific cod (TL 4.25) has a

632 predicted concentration of 114.3 Bq·t-1. In the comparison between these two species the

633 biggest differences are between the Q/B and the P/B ratio. The Q/B rate is 65 percent higher

634 for Alaska pollock, and the P/B ratio is 21 percent lower indicating a lower turnover rate

635 implying that this group has a longer time to accumulate 137Cs in their tissues.

636
 28

637 The 137Cs concentration generally biomagnifies throughout the ecosystem with a mean value

638 of 1.92. This is in agreement with laboratory feeding experiments (Wang et al. 2000; Zhao et

639 al. 2001; Matthews and Fisher 2008), field measurements (Kasamatsu and Ishikawa 1997;

640 Harmelin-Vivien et al. 2012), and in a food web model (Alava and Gobas 2016). However, 4

641 invertebrate groups were well below a BMF of 1 indicating that in these groups 137Cs did not

642 biomagnify. Of these, herbivorous zooplankton were basically equal to their main diet item

643 of copepods, but the other three groups (squid, crabs & lobsters, and seastars) were below

644 0.5. For the forage fish that primarily eat copepods, physiological processes may limit the

645 accumulation of 137Cs since this groups had the highest predicted elimination rate constant

646 (Ke) of all fish groups (Supplementary data 2). For the 3 other fish groups with BMF values

647 greater than 0.9, uncertainty surrounding the diet matrix or physiological processes could

648 explain why 137Cs does not biomagnify.

649

650 For the groups in the area with reported concentrations of 137Cs, the model did well in

651 replicating the measured values.. The model was also able to make predictions for groups

652 without initial concentrations that, based on the comparison of CRs for groups with starting

653 concentration data and those without starting concentrations (Figure 2), would seem to be

654 within expected ranges. The importance of setting a baseline in this model is for future

655 scenarios that do not rely on the CR approach where the resulting concentrations are driven

656 by changes in the environmental concentrations only. In these scenarios, the use of a

657 hydrodynamic model would be useful to explain the 137Cs water concentrations and the

658 resulting 137Cs concentrations in biota.

659
 29

660 The groups below or above the trend line for biomass in the PREBAL diagnostic represent a

661 trade-off between the Ecopath model structure and the Ecotracer function of tracing 137Cs

662 through the ecosystem. Here, it was decided to define groups on the basis of the

663 information available regarding 137Cs concentrations. Keeping groups with relatively good

664 data regarding 137Cs concentrations separate, rather than placing them within larger

665 functional groups, allows the concentration data to be retained and to be used for

666 performance tests.

667

668 Recommendations for the improvement of the Ecotracer routine within the EwE ecological

669 modelling framework include: 1) separating the environment into depth zones; 2)

670 implementing the separation between metabolic and excretory pathways; and, 3) including a

671 routine for dose assessment.

672

673 Many pollutants have a vertical profile of concentration that either increases or decreases

674 with depth. Dividing the environment into depth zones would make the Ecotracer routine

675 adaptable to a specific contaminant’s water column profile. Depth zones could include

676 coastal, mid-water and deep-water zones, and could be linked to the habitat capacity model

677 within Ecospace that already includes the ability to assign functional groups to depth zones.

678

679 In the latest version of Ecotracer, it is not possible to differentiate between metabolic losses

680 and excretory losses as described in Walters and Christensen (2018). Metabolic losses occur

681 when the contaminant undergoes a transformation and thus is lost from the system,

682 whereas when it is excreted from the organism’s body it remains as part of the

683 environmental pool. Currently, the best option is to include both as metabolic losses, but an
 30

684 error is introduced as the portion included from excretory losses are removed from the

685 environment rather than being in the environment and being available for direct uptake.

686

687 Estimated concentrations in seafood were all well below the Japanese pre-accident

688 regulatory limit of 500 Bq·kg-1. The apparent supply of 137Cs represents the amount in the

689 fishery products before processing and includes portions that may not be eaten. However, it

690 can help to determine the health effects on the human body from the consumption of

691 contaminated seafood. In this respect, Ecotracer could also be improved to allow dose

692 assessment for both marine organisms and humans, and made valuable to health agencies

693 and risk assessors. For marine organisms, a special routine for assessing organism dose rates

694 arising both from internal and external exposures would need to be developed. In terms of

695 protecting human health, effective dose from ingestion can be estimated through the use of

696 dose conversion coefficients (International Commission on Radiological Protection 2012).

697

698

699 ACKNOWLEDGEMENTS

700

701 This work has been funded by the Institute de Radioprotection et de Sûreté Nucléaire and by

702 the French program Investissement d’Avenir run by the National Research Agency (AMORAD

703 project, grant ANR-11-RSNR-0002). The authors would like to thank the Central Research

704 Institute of Electric Power Industry (Abiko, Japan) for hosting SB for one month and Yutaka

705 Tateda for assisting with the translation of articles in Japanese. The authors would also like

706 to thank Japan’s Ministry of Agriculture Forestry and Fishery for providing fisheries data
 31

707 when requested, two anonymous reviewers, and the editor for enhancing the manuscript

708 through their comments.

709

710

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Figure 1: Flow diagram representing the trophic structure of the marine ecosystem of Iwate, Miyagi, Fukushima, and Ibaraki prefectures of the Pacific coast
of eastern Japan in 2010. Fleet 1 represents all the fishing fleets operating within the exclusive economic zone for the prefectures of Iwate, Miyagi,
Fukushima and Ibaraki.
Figure 2: Distribution of concentration ratios from the EwE model for groups with data (observed),
and those groups that lacked data reported in the literature (predicted). Median values were 44.9
and 43.6 for observed and predicted values, respectively. Including the outlier for the observed value
increases the median value to 47.0.
Figure 3: Observed activity (Bq) versus predicted activity (Bq) for the concentration of 137Cs in the
groups with concentration ratios reported in the literature.
Figure 4: increase of 137Cs by trophic level in the marine ecosystem of the four prefectures (Iwate,
Miyagi, Fukushima and Ibaraki) of eastern Japan. Solid circles represent those groups in the model
with CRs, hollow circles are groups lacking data and are estimated by Ecotracer.
Table 1. Biomass (t∙km-2), production to biomass ratio, (P/B; year-1); consumption to biomass ration (Q:B
;year-1); ecotrophic efficiency (EE), trophic level (TL), production to consumption ratio (P/Q) and fishery
landings (t∙km-2∙year-1) for the Ecopath model for the marine ecosystem model of Iwate, Miyagi,
Fukushima and Ibaraki prefectures representing about 220,000 km-2. EwE model estimates are in bold.
No. Group Biomass P/B Q/B EE TL P/Q Landings
1 Large sharks 0.080 0.389 1.60 0.683 4.77 0.243 0.01870
2 Small benthivorous sharks 0.060 0.946 3.83 0.783 3.87 0.247 0.03740
3 Small piscivorous sharks 0.060 0.818 3.85 0.942 4.62 0.212 0.03740
4 Skates & rays 0.033 0.510 2.53 0.950 4.23 0.202 0.01250
5 Large pelagics 0.040 0.970 3.60 0.950 4.78 0.269 0.03732
6 Medium pelagics 0.436 0.760 6.50 0.950 4.48 0.117 0.29612
7 Small pelagics 0.129 0.910 6.60 0.950 4.58 0.138 0.00540
8 Deep benthopelagics 0.052 0.420 2.79 0.950 4.32 0.151 0.01250
9 Shallow benthopelagics 0.095 0.840 4.90 0.950 4.13 0.171 0.04611
10 Halibut 0.017 0.420 4.40 0.878 4.03 0.095 0.00104
11 Pacific cod 0.525 0.484 2.30 0.599 4.25 0.210 0.10810
12 Fat greenling 0.063 0.742 4.40 0.950 4.00 0.169 0.01250
13 Omivorous bathydemersals 1.023 0.240 2.37 0.950 4.10 0.101 0.01250
14 Crustivorous bathydemersals 0.311 0.220 2.15 0.950 3.76 0.102 0.01870
15 Alaska pollock 1.710 0.380 3.80 0.756 3.74 0.100 0.07412
16 Bathy-meso pelagics 1.825 1.500 9.00 0.950 3.38 0.167 -
17 Pelagic neritic piscivores 0.009 1.070 7.10 0.950 4.60 0.151 0.00630
18 Pelagic neritic planktivores 2.262 1.311 7.40 0.950 3.52 0.177 0.29300
19 Reef piscivores 0.130 1.002 6.68 0.950 4.27 0.150 0.01250
20 Reef benthivores 0.885 0.725 4.83 0.950 3.50 0.150 0.01250
21 Reef planktivores 0.721 1.140 7.60 0.950 3.59 0.150 0.00091
22 Sebastes 0.360 0.654 4.36 0.950 3.74 0.150 0.01250
23 Demersal-continental slope 0.055 0.526 2.63 0.950 4.10 0.200 0.00630
24 Demersal-rocky 0.028 0.515 3.33 0.859 4.16 0.155 0.00300
25 Demersal-sandy/mud 1.984 0.899 6.56 0.950 3.56 0.137 0.05748
26 Pufferfish 0.139 1.050 5.26 0.950 3.32 0.200 0.01250
27 Deep flounders 0.026 0.420 2.40 0.923 4.07 0.175 0.00357
28 Shallow flounders 0.128 0.560 5.24 0.931 3.56 0.107 0.00712
29 Deep soles 0.020 0.420 2.30 0.943 3.55 0.183 0.00357
30 Shallow soles 0.149 0.560 3.73 0.790 3.37 0.150 0.00712
31 Olive flounder 0.020 0.560 2.40 0.950 4.39 0.233 0.00418
32 Flathead flounder 0.026 0.420 4.20 0.950 3.74 0.100 0.00209
33 Marbled flounder 0.050 0.560 4.00 0.950 3.38 0.140 0.00209
34 Mackerels 2.350 0.777 7.70 0.742 3.63 0.101 0.66356
35 Japanese jack mackerel 0.070 0.944 3.60 0.850 3.07 0.262 0.02053
36 Forage fish-euphausivores 3.337 1.176 6.77 0.950 3.44 0.174 0.39070
37 Forage fish-copepodivores 0.122 1.180 10.23 0.927 3.01 0.115 0.00120
38 Octopus 0.227 1.200 10.60 0.950 3.68 0.113 0.02655
39 Crabs & lobster 7.513 0.800 5.00 0.950 3.01 0.160 0.00368
40 Squids 2.583 3.080 10.80 0.950 3.87 0.285 0.18415
41 Seastars 1.708 1.220 5.00 0.950 2.65 0.244 -
42 Mantis shrimp 0.723 0.912 4.56 0.950 3.48 0.200 0.00004
43 Omnivorous shrimp 7.656 1.440 7.20 0.950 2.61 0.200 0.00031
44 Sea cucumbers 1.569 0.500 5.00 0.950 2.00 0.100 0.00091
45 Sea urchins 0.453 0.610 5.00 0.950 2.00 0.122 0.01049
46 Annelids 7.100 2.930 12.00 0.950 2.36 0.244 -
47 Gastropods 6.803 1.810 8.00 0.950 2.39 0.226 -
48 Bivalves 15.637 1.470 12.00 0.950 2.00 0.123 0.01718
49 Krill 11.990 11.120 44.50 0.592 2.50 0.250 0.18780
50 Carnivorous zooplankton 20.700 4.560 15.00 0.627 3.04 0.304 -
51 Omnivorous zooplankton 2.710 4.880 16.30 0.893 2.86 0.299 -
52 Herbivorous zooplankton 101.00 5.200 20.00 0.965 2.00 0.260 -
53 Detritivores 25.153 3.530 17.65 0.950 2.00 0.200 -
54 Macroalgae 0.834 16.000 - 0.950 1.00 - 0.01136
55 Phytoplankton 12.000 190.000 - 0.61 1.00 - -
56 Detritus 146.00 - - 0.643 1.00 - -
Table 2. Ecotracer initial concentrations (Bq·t-1), direct absorption rates (DAR,ui; km2∙t-1∙year-1), physical decay rates (PDR;
year-1), fraction excreted (1-AE), excretion rates (Ke; year-1), final concentrations (Bq·t-1), proportion of final concentration
due to respiration (Presp), and the biomagnification factor (BMF).
No. Group [137Cs]i DAR (ui) PDR 1-AE Ke [137Cs]f Presp BMF
1 Large sharks 0.00 1.07 x 10-4 0.023 0.200 0.889 108.1 0.384 1.60
2 Small benthivorous sharks 0.00 4.07 x 10-4 0.023 0.200 3.257 72.9 0.664 2.14
3 Small piscivorous sharks 0.00 7.77 x 10-5 0.023 0.200 1.134 100.3 0.197 1.95
4 Sharks & rays 196.80 7.32 x 10-4 0.023 0.200 1.999 197.5 0.742 3.10
5 Large pelagics 0.00 1.95 x 10-4 0.023 0.375 0.867 109.6 0.485 3.08
6 Medium pelagics 0.00 0.0 0.023 0.375 1.404 94.1 0.000 1.88
7 Small pelagics 0.00 0.0 0.023 0.200 1.176 99.1 0.000 1.95
8 Deep benthopelagics 76.50 7.34 x 10-5 0.023 0.200 1.697 76.4 0.226 1.41
9 Shallow benthopelagics 94.50 1.97 x 10-4 0.023 0.200 2.207 93.5 0.342 2.14
10 Halibut 75.00 1.69 x 10-4 0.023 0.200 2.614 75.0 0.372 1.85
11 Pacific cod 114.00 2.42 x 10-4 0.023 0.200 1.943 113.8 0.438 1.34
12 Fat greenling 67.50 1.13 x 10-4 0.023 0.200 2.306 65.8 0.274 1.54
13 Omnivorous bathydemersals 48.75 6.38 x 10-5 0.023 0.200 2.374 48.6 0.251 0.95
14 Crustivorous bathydemersals 0.00 3.68 x 10-4 0.023 0.200 3.874 69.5 0.648 1.17
15 Alaska pollock 141.00 9.15 x 10-4 0.023 0.200 3.920 141.0 0.758 2.91
16 Bathy-meso pelagics 0.00 5.46 x 10-4 0.023 0.200 6.674 59.0 0.570 1.60
17 Pelagic neritic piscivores 0.00 0.0 0.023 0.546 1.165 98.3 0.000 1.42
18 Pelagic neritic planktovores 0.00 1.12 x 10-4 0.023 0.200 5.513 62.6 0.133 0.98
19 Reef piscivores 90.00 0.0 0.023 0.422 1.784 88.6 0.000 1.34
20 Reef benthivores 0.00 4.74 x 10-4 0.023 0.200 5.528 62.1 0.611 1.54
21 Reef planktivores 0.00 1.89 x 10-4 0.023 0.200 5.309 63.3 0.234 1.21
22 Sebastes 99.90 5.29 x 10-4 0.023 0.200 3.907 99.8 0.583 1.78
23 Demersal-continental slope 0.00 2.33 x 10-4 0.023 0.200 2.316 80.9 0.506 1.40
24 Demersal-rocky 73.50 6.56 x 10-5 0.023 0.200 2.187 73.2 0.165 1.17
25 Demersal-sandy/mud 98.25 7.13 x 10-4 0.023 0.200 5.519 97.9 0.603 2.12
26 Pufferfish 61.50 6.61 x 10-4 0.023 0.200 7.209 61.4 0.655 1.43
27 Deep flounders 0.00 2.50 x 10-4 0.023 0.200 2.498 79.3 0.541 1.42
28 Shallow flounders 58.80 3.02 x 10-4 0.023 0.200 5.139 58.6 0.454 1.32
29 Deep soles 0.00 6.15 x 10-4 0.023 0.200 5.225 63.5 0.861 2.29
30 Shallow soles 0.00 5.97 x 10-4 0.023 0.200 6.765 58.7 0.697 1.31
31 Olive flounder 103.50 1.02 x 10-4 0.023 0.200 1.601 101.8 0.227 1.11
32 Flathead flounder 63.00 2.65 x 10-4 0.023 0.200 4.008 62.6 0.478 1.42
33 Marbled flounder 97.50 1.15 x 10-4 0.023 0.200 6.617 71.9 0.829 2.30
34 Mackerels 0.00 0.0 0.023 0.219 5.584 62.8 0.000 0.93
35 Japanese jack mackerel 0.00 1.05 x 10-3 0.023 0.200 10.293 51.8 0.910 2.83
36 Forage fish-euphausivores 117.0 1.13 x 10-3 0.023 0.200 6.143 117.0 0.665 2.21
37 Forage fish-copepodivores 0.00 4.59 x 10-4 0.023 0.200 11.204 34.6 0.370 0.70
38 Octopus 31.50 6.77 x 10-5 0.023 0.500 4.256 31.5 0.198 1.20
39 Crabs & lobsters 15.00 1.58 x 10-4 0.023 0.500 10.537 15.0 0.469 0.40
40 Squids 13.50 0.0 0.023 0.898 2.927 13.5 0.000 0.18
41 Seastars 0.00 3.03 x 10-4 0.023 0.500 18.506 10.5 0.738 0.47
42 Mantis shrimp 0.00 2.27 x 10-3 0.023 0.300 10.950 108.0 0.893 2.43
43 Omnivorous shrimp 0.00 4.89 x 10-3 0.023 0.350 41.975 60.0 0.948 2.02
44 Sea cucumbers 0.00 3.13 x 10-3 0.023 0.350 47.450 33.0 0.999 5.16
45 Sea urchins 0.00 2.45 x 10-3 0.023 0.500 38.690 33.0 0.955 1.35
46 Annelids 0.00 1.27 x 10-2 0.023 0.840 162.425 39.0 0.995 2.44
47 Gastropods 55.50 3.03 x 10-3 0.023 0.500 27.010 55.5 0.956 3.06
48 Bivalves 25.50 3.31 x 10-3 0.023 0.835 64.970 25.4 0.987 3.47
49 Krill 60.00 5.56 x 10-3 0.023 0.350 41.975 60.0 0.880 4.89
50 Carnivorous zooplankton 108.00 2.79 x 10-3 0.023 0.300 10.959 108.0 0.840 4.45
51 Omnivorous zooplankton 60.00 1.59 x 10-3 0.023 0.350 13.848 60.1 0.712 1.94
52 Herbivorous zooplankton 12.00 9.07 x 10-4 0.023 0.350 46.059 12.1 0.744 0.99
53 Detritivores 33.0 3.19 x 10-3 0.023 0.350 47.450 33.1 0.956 5.20
54 Macroalgae 27.45 1.08 x 10-2 0.023 0.000 182.500 27.5 1.000 -
55 Phytoplankton 12.75 9.41 x 10-3 0.023 0.000 182.500 12.7 1.000 -
56 Detritus 6.30 - 0.023 - - 6.6 - -