Geotrupine beetles (Coleoptera: Scarabaeoidea) as bio-monitors of

man-made radioactivity

Jerzy W. Mietelski,*a Przemysław Szwałko,b Ewa Tomankiewicz,a Paweł Gacaa and

Sylwia Grabowskaa

aThe Henryk Niewodniczanski Institute of Nuclear Physics, Krakow, Poland.E-mail: mietelsk@alf.ifj.edu.pl; Fax: 148 12 6628458; Tel: 148 12 6628 392

bInstitute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow,Poland

Received 3rd January 2003, Accepted 10th February 2003

First published as an Advance Article on the web 20th February 2003

Adults of the geotrupine beetle Anoplotrupes stercorosus (Coleoptera, Geotrupidae), a common European forest

insect species, were used in the role of bio-monitors for mainly man-made radionuclides in a forest

environment. Activities of 137Cs, 40K, 238Pu, 2391240Pu, 90Sr and 241Am were studied. Samples originated from

four areas in Poland, two from the north-east and two from the south of the country. The north-eastern areas

were previously recognized as the places where hot particle fallout from Chernobyl took place. Results

confirmed the differences in the activities between north-eastern and southern locations. Significant correlations

were found between activities of 40K and 137Cs, and between activities of plutonium and americium isotopes.

An additional study of the concentration of radionuclides within the bodies of beetles showed a general pattern

of distribution of radioisotopes in the insect body.

Introduction

The beetle Anoplotrupes stercorosus (Coleoptera, Geotrupidae)is one of the most common and relatively large European forestinsects. The imago individual is about 2 cm long with a freshmass between 0.30 g and 0.45 g. It lives in the forests of almostthe whole Palaearctic boreal zone. The adults usually feed onfungi or dung, and can also be attracted to carrion. The larvalfood contains mainly dead leaves and humus, occasionallymixed with other organic matter as particles of deer dung ormushroom fruit bodies. The larva develops ca. 20 cm deepin the soil, taking nutrients from humus clusters formed thereby its parents. Development depends on climatic conditionsand requires one or two years. Young imagines appear usuallyin late summer and survive winter in hibernation until thenext vegetation season. Geotrupine beetles can be collected asimago using unbaited pitfall traps in large quantities. For anumber of years this species was used for bio-monitoring.1–4

The humus layer of forest litter contains a lot of man-made radionuclides, namely 137Cs, 2391240Pu, 238Pu, 241Am, andsome 90Sr present still there from global fallout which tookplace mostly in the fifties and sixties of the twenty century aswell as some newer radionuclides, from the Chernobyl fallout.In north-eastern Poland, Chernobyl hot particle falloutoccurred, resulting (in 1986) in deposited activities of up to30 Bq m22 of Pu alpha isotopes,5 another6 1 kBq m22 of241Pu and about7,8 2 kBq m22 of 90Sr, whereas 137Cs waspresent on levels rarely exceeding9 5 kBq m22. Maximumvalues were found for the Augustow Primeval Forest area.The south of Poland had a typical isotopic composition ofChernobyl fallout for the remote sites, with 137Cs deposition9

ranging from a few to 50 kBq m22 and with almost exclusiveglobal fallout origin of 90Sr, plutonium or americium activ-ities. The other difference between those two areas is that thesouth of Poland is far more contaminated with all kindsof industrial pollution than north-eastern Poland, which isconsidered as the ‘‘green lungs of Poland’’.

Material and method

Samples of Anoplotrupes stercorosus (Hartm.) were collectedbetween 1989 to 2001 from two huge forest complexes ofnorth-eastern Poland, mainly from Białowieza PrimevalForest (63 150 ha) but also from Augustow Primeval Forest(101 669 ha), and at two control locations in southern Poland,mainly from Niepołomice Primeval Forest (11 000 ha) but alsofrom Tułowice Forest (9990 ha) in Silesia. The Tułowicelocation is the site situated near the area with the highest 137Cspost-Chernobyl contamination in Poland, at the level9 of30 kBq m22. Augustow forest is the location with the mostintensive hot particle fallout (Fig. 1). Some data on samplinglocations are presented in Table 1. For all sampling areas the

Fig. 1 Approximate map of Chernobyl-origin 2391240Pu deposition inPoland,18 based on 144Ce measurements in forest litter.19 Dots aresampling sites for 144Ce measurements. Locations of sampling sites forthe present study: A—Augustow, B—Białowieza, N—Niepołomice,T—Tułowice.

296 J. Environ. Monit., 2003, 5, 296–301 DOI: 10.1039/b212592g

This journal is # The Royal Society of Chemistry 2003

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pH of soil surface on A horizon (under forest litter) was alwaysacidic, ranging from 4.7 (for fresh deciduous forest) to 3.8(for fresh mixed coniferous or wet deciduous forests).10 Twoadditional samples of Trypocopris vernalis (L.) were alsoanalyzed for comparison. This is another representative of theGeotrupine beetle family, and it looks similar to A. stercorosus(Fig. 2). However, its feeding habits are different. It does noteat humus, but it is a coprophag, which feeds on faeces ofherbivorous mammals both as larva and as imago. One sampleof Trypocopris vernalis was trapped by chance together with A.stercorosus in Białowieza (collected during 1990–99), the othersample was similar but from the Tułowice area.All samples were analyzed for the presence of radionuclides

taking whole bodies of adult insects (imagines). Samplescontained from 67 to 750 imagines of beetle, with the arithmeticmean equal to 317 and the standard deviation of 174. Analyseswere performed between summer of 2001 and winter of 2001/2002 and the measurements lasted till the summer of 2002.Besides measurements of whole beetles, five kind of organs

were separated out of 180 beetle bodies from Augustow(collected in 2001). The separated organs were: heads, legs(mainly skeleton and muscles), elytra (wing covers, a prettypure skeleton), internal organs of thorax and abdomen(including genital organs and digesting duct), and remainingbody parts. This study was done with the aim of getting an ideaof the distribution of radionuclides within the beetle body.For each one whole body sample insects were counted

and initial mass was determined. Next, samples were dried anddry masses were determined, then incinerated at 400 uC in theoven and the ash masses were again determined. Some samplesoriginated from past collections, remaining from formerprojects,2–4 some others were newly collected in 2001. Someof the archive samples were air dried, some were kept in ethanol(alcohol was also taken for analyses, there were no radio-nuclides found). Because of those differences we were unable touse the concepts of both fresh or dried masses of animals. Wedecided to present in this paper the results normalized for themass of ashes of animal incinerated at 400 uC. As the average

beetle has a fresh mass in the range of about 0.30–0.45 g, themean (¡ standard deviation) mass of ashed beetle was0.0023 ¡ 0.0013 g.Ashes were soaked with a water solution containing 0.4 g of

glucose and evaporated to dryness to make a solidified tablet.The activities of 137Cs and 40K (natural isotope) in ashes weremeasured by means of low background gamma-ray spectro-metry with a coaxial HPGe detector, in standard for us samplegeometry of a shallow cylinder (50 mm diameter). Selectedtissues/organs of beetle were measured inside the well of awell-type HPGe detector calibrated with 0.5 g of IAEA 375Reference Material.11

After these measurements the radioactive tracers (85Srproduced by INP, 236Pu produced by Harwell and 243Am byNBS) were added and all the samples were mineralized bysubsequent adding and then boiling until evaporation to almostdryness of following portions of reagents: 20 ml of 20 M HF,2 6 20 ml of 14 M HNO3, 20 ml of 14 M HNO3 with 5 ml ofHClO4, 5 ml H2O2, 30 ml of 12 M HCl with 1 g of H3BO3

added and finally 25 ml of 14 M HNO3. The last solution wasevaporated only to about 10 ml and then diluted with water toabout 1 M HNO3. Perchloric acid and peroxide were added todestroy any traces of glucose or carbon. After this treatment,samples were colorless and nicely transparent, but they werefiltered just to avoid any possible problems with any suspendedmatter during the following steps. The Pu oxidation adjustmentprocedure,12 using hydrazine and NaNO2 was applied. Radio-nuclides were separated sequentially, in the course of onecombined procedure. Pu was separated12 on an anion exchangecolumn filled with Dowex-1 from 8 M HNO3, Sr on

13 Sr-Resinalso from 8 M HNO3, and finally Am was separated usingDowex-1 and methanol–acid solutions.14 Alpha spectrometricsources (Pu, Am) were prepared using NdF3 co-precipitationmethod.15,12,5 The measurements were performed using aSilena AlphaQuattro spectrometer with Canberra PIPSdetectors. The activity of 90Sr was determined using a liquidscintillation spectrometer Wallac 1414-003 Guardian, with the85Sr recovery determined by means of gamma spectrometry.

Results and discussion

The activities of 137Cs, 40K, 2391240Pu, 238Pu, 90Sr, and 241Am,normalized for ash weight of samples are presented in Table 2.Since the individual sample consists of a large number ofindividual insects, the obtained results might be consideredclose to the mean values for the population. What is perhaps alittle striking is that the activities at given areas did not changesmoothly from year to year, but were a bit more scattered thanone might expect. The mean recoveries of tracers and theirstandard deviations were (76 ¡ 20)%, (74 ¡ 28)% and (42 ¡13), for plutonium, strontium, and americium, respectively.For 137Cs, Pu, Am, and 90Sr the highest activities in ashes

of A. stercorosus for samples from north-eastern Poland wereobserved for a sample subdivided into two halves, collected in

Table 1 Data on sampling areas

Approximate position of the area Sample siteYear ofsampling Forest site type

137Cs in surface soil9/kBq m22

Białowieza primeval forest 53u00’N, 23u32’E b3 1993, 1997 Fresh deciduous forest v3 to 15b5 1990, 1992–99 Fresh mixed coniferous forest

Augustow primeval forest 54u04’N, 23u05’E a/89 1989 Fresh mixed coniferous forest v3 to 8a/01 2001 Fresh mixed coniferous forest

Niepołomice primeval forest 50u02’N, 20u22’E n1 1993, 1994 Fresh mixed deciduous forest v3 to 8n2 1993, 1994 Fresh mixed coniferous forestn3 1993, 1994 Weta mixed deciduous forestn5 1993, 1994 Weta deciduous forest

Tułowice forest 50u40’N, 17u32’E t 1994–96 Fresh coniferous forest y30t 2001 Fresh mixed coniferous forest

aMore humid than ‘‘fresh’’.

Fig. 2 Geotrupine beetles: (a) Anoplotrupes stercorosus (Hartm.), (b)Trypocopris vernalis (L.).

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Augustow forest in 2001. The maximum values were equal to6.72 ¡ 0.56 kBq kg21, 5.7 ¡ 0.6 Bq kg21, 1.5 ¡ 0.2 Bq kg21,3.0 ¡ 0.6 Bq kg21, 0.16 ¡ 0.02 Bq g21 for 137Cs, 2391240Pu,238Pu, 241Am, and 90Sr, respectively. Moreover, the traces of2441243Cm (at the level of 0.15 ¡ 0.09 Bq kg21 on average)were found in this sample as well.The highest activities in ashes for samples from southern

Poland were found for various radionuclides in differentsamples. The maximum value for 137Cs was found for samplesfrom Tułowice forest, and was equal to 13.8 ¡ 0.2 Bq kg21.This follows expectations since the area has a high 137Csdeposition. A noticeable difference was found between sampl-ing sites in Niepołomice forest. The samples from sites n2 andn3 have an order of magnitude higher activity of 137Cs thanthose from site n5, whereas samples from site n1 haveintermediate values. This is not the case for the Pu or Amisotopes. The main factor for this difference are differences inthe forest type. The n2 site is mixed coniferous, whereas n1 andn3 sites are mixed deciduous and n5 is deciduous forest. On theother hand, a spot-like, or fractal pattern of Chernobyl fallout,which was the source of the majority of 137Cs activity (butnot Pu nor Am isotopes) in southern locations, may influenceobserved variety.It seems that activities found for T. vernalis are lower than

those for A. stercorosus.Data now obtained for Anoplotrupes stercorosus confirmed

the differences between isotopic composition of Chernobylfallout in north-eastern (NE) Poland and in the south (S) of thecountry. The influence of Chernobyl hot particle depositionwas visible and distinguishable, especially for the sample fromAugustow primeval forest. Activities of 2391240Pu, 241Am, andespecially 238Pu were significantly higher for NE samples. Interms of the t-test with Welch correction the difference in

activities of 238Pu or 241Am between NE locations (Białowieza,Augustow) and S ones (Niepołomice, Tułowice) are considered‘‘extremely significant’’, with the p value equal to 0.0006 in bothcases. The Mann–Whitney test also recognizes the difference as‘‘extremely significant’’. For 2391240Pu the difference betweensouthern and north-eastern Poland is ‘‘very significant’’ interms of both tests mentioned above. However, the differencefor 137Cs is considered ‘‘not significant’’.The observed differences between the radionuclides content

in examined beetles from the area with hot particle depositionand the area without it, suggest that isotopes present in hotparticles were previously already bio-available during thesampling. In Fig. 3 the time dependence of 238Pu to 239Puactivity ratio for samples are presented. The global falloutvalue16 for this ratio is 0.03, whereas for the Chernobyl falloutin Poland5 it was about 0.55. The average value (and standarddeviation) for NE samples was 0.203 ¡ 0.083. It means about33% of 2391240Pu was of Chernobyl origin. The same ratio forS samples is equal to 0.135 ¡ 0.154. This gives about 20% ofChernobyl-origin Pu. However, if one would remove only onepossible outlying result which has relatively large uncertainty,the average drops to 0.097 ¡ 0.102 resulting in 13% ofChernobyl Pu. In Fig. 4 the activity ratio for 241Am to 239Pu isdisplayed. The average values were equal to 0.451¡ 0.189 and0.360 ¡ 0.220, for NE and S samples, respectively. Again theyare higher for NE areas then for S, where the average is closeto the global fallout value for this ratio equal16 to about 0.3.The present Chernobyl ratio17 (including 241Am ingrown from241Pu decay) is about 1.5, so it is a much higher value thanfound in NE samples. The difference in activities ratios betweensamples from S and NE location, as discussed here, can be seen,but only a possible very weak activity ratio increasing with timetendency for 241Am to 2391240Pu was suggested for samples

Table 2 Results for activities found in ashen beetle imagines Anoplotrupus stercorosus and Trypocopris vernalis

Code/site

137Cs/Bq(g-ash wt.)21

40K/Bq(g-ash wt.)21

2391240Pu/Bq(kg-ash wt.)21

238Pu/Bq(kg-ash wt.)21

90Sr/Bq(g-ash wt.)21

241Am/Bq(kg-ash wt.)21

A. stercorosusBiałowiezab3/94 0.51 ¡ 0.02 1.91 ¡ 0.15 0.52 ¡ 0.08 0.04 ¡ 0.03 v0.010 0.24 ¡ 0.08b3/97 0.49 ¡ 0.02 1.31 ¡ 0.17 3.24 ¡ 0.28 0.40 ¡ 0.13 0.013 ¡ 0.003 0.39 ¡ 0.21b5/90 5.53 ¡ 0.11 2.37 ¡ 0.52 6.03 ¡ 0.45 1.17 ¡ 0.24 v0.036 1.00 ¡ 0.45b5/92 1.74 ¡ 0.04 5.24 ¡ 0.32 2.47 ¡ 0.20 0.77 ¡ 0.15 v0.025 1.30 ¡ 0.40b5/93 1.98 ¡ 0.09 1.40 ¡ 0.67 5.85 ¡ 0.46 0.95 ¡ 0.21 No data 2.92 ¡ 0.76b5/94 1.19 ¡ 0.05 3.14 ¡ 0.43 3.64 ¡ 0.25 1.34 ¡ 0.46 v0.056 0.87 ¡ 0.81b5/95 2.37 ¡ 0.07 2.29 ¡ 0.33 1.07 ¡ 0.10 0.09 ¡ 0.06 No data 0.89 ¡ 0.52b5/96 1.94 ¡ 0.05 2.63 ¡ 0.29 2.26 ¡ 0.20 0.64 ¡ 0.08 v0.023 1.24 ¡ 0.31b5/97 1.23 ¡ 0.06 1.38 ¡ 0.38 3.40 ¡ 0.32 0.55 ¡ 0.13 0.028 ¡ 0.040 1.80 ¡ 0.67b5/98 1.59 ¡ 0.04 1.89 ¡ 0.23 3.06 ¡ 0.24 0.67 ¡ 0.13 0.013 ¡ 0.004 1.72 ¡ 0.41b5/99 0.95 ¡ 0.03 1.39 ¡ 0.23 1.88 ¡ 0.16 0.37 ¡ 0.08 No data 1.09 ¡ 0.44Augustowa/89 3.20 ¡ 0.15 4.25 ¡ 1.16 4.44 ¡ 0.50 0.80 ¡ 0.25 No data 1.53 ¡ 0.75a/01–1 6.72 ¡ 0.08 6.91 ¡ 0.47 5.49 ¡ 0.39 1.49 ¡ 0.17 0.142 ¡ 0.018 3.01 ¡ 0.50a

a/01–2 6.71 ¡ 0.06 8.48 ¡ 0.36 5.69 ¡ 0.40 1.20 ¡ 0.13 0.161 ¡ 0.021 2.00 ¡ 0.22a

Niepołomicen1/93 1.68 ¡ 0.06 4.58 ¡ 0.55 1.83 ¡ 0.27 0.18 ¡ 0.16 0.025 ¡ 0.007 0.62 ¡ 0.21n1/94 0.96 ¡ 0.05 3.09 ¡ 0.54 0.71 ¡ 0.08 0.01 ¡ 2.27 0.029 ¡ 0.006 0.08 ¡ 0.08n2/93 5.72 ¡ 0.11 5.24 ¡ 0.70 2.90 ¡ 0.45 0.02 ¡ 0.56 0.019 ¡ 0.005 0.67 ¡ 0.40n2/94 3.85 ¡ 0.24 4.75 ¡ 2.16 4.14 ¡ 1.04 0.41 ¡ 0.33 0.050 ¡ 0.025 0.66 ¡ 0.23n3/93 5.37 ¡ 0.18 6.67 ¡ 1.31 1.83 ¡ 0.48 0.14 ¡ 0.28 0.055 ¡ 0.037 0.72 ¡ 0.48n3/94 3.53 ¡ 0.11 4.95 ¡ 0.97 1.19 ¡ 0.40 0.02 ¡ 0.07 0.049 ¡ 0.018 0.81 ¡ 0.41n5/93 0.24 ¡ 0.02 3.11 ¡ 0.36 1.04 ¡ 0.19 0.50 ¡ 0.20 0.048 ¡ 0.013 0.21 ¡ 0.07n5/94 0.31 ¡ 0.05 4.67 ¡ 0.65 0.40 ¡ 0.11 0.09 ¡ 0.08 0.055 ¡ 0.017 0.32 ¡ 0.15Tułowicet/94–96 7.82 ¡ 0.15 4.66 ¡ 0.88 1.54 ¡ 0.18 0.46 ¡ 0.10 0.041 ¡ 0.007 0.51 ¡ 0.14t/01 13.78 ¡ 0.21 9.45 ¡ 1.54 1.14 ¡ 0.20 0.04 ¡ 0.10 v0.027 0.41 ¡ 0.25T. vernalisBiałowiezatb/90–99 1.22 ¡ 0.07 1.99 ¡ 0.90 1.71 ¡ 0.21 0.43 ¡ 0.12 0.181 ¡ 0.055 0.22 ¡ 0.11Tułowicett/94–95 6.95 ¡ 0.14 4.73 ¡ 1.01 0.31 ¡ 0.09 0.31 ¡ 0.09 0.032 ¡ 0.008 0.23 ¡ 0.09a244Cm also found.

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from Białowieza forest. It should be pointed out that thispossible tendency rather has nothing to do with the obviousingrown 241Am from 241Pu decay, since all the samples wereanalyzed recently, during the course of only one year, so 241Ambuild-up from 241Pu is at a similar level for all samples. Thismight suggest a little higher transfer factor for Am than for Pufrom litter to beetle.

In Fig. 5 the most important correlations between observedactivities are displayed. The activities of both plutoniumisotopes were significantly correlated to each other, and with241Am activity, but not with 137Cs. The last one correlatedmostly with 40K.The results on radionuclide distribution in the insect body are

presented in Tables 3 and 4. About 45% of each radionuclide

Fig. 3 The variety of 238Pu to 2391240Pu activity ratio for samples collected in different years in north-eastern Poland (left) and southern Poland(right).

Fig. 4 The time variety of 241Am to 2391240Pu activity ratio for samples collected in different years in north-eastern Poland (left) and southernPoland (right). A weak tendency (straight lines fit) is suggested for north-eastern samples.

Fig. 5 Correlation plots for activities found in Anoplotrupes stercorosus samples. 2391240Pu correlates significantly with 241Am and 238Pu, and 137Cscorrelates with 40K, but not with 2391240Pu. R is Pearson’s correlation factor, p is the level of significance.

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activity was found in ‘‘remaining body parts’’ (Table 3). About35% of 90Sr was found in legs. Plutonium and americiumwas divided mainly between some separated internal organs(containing the digestion system) and remains of the body (withsome other inner organs left). The figure became more clearafter normalization of the activity to the percentage of the massof the body ashes which a given, selected organ has. Suchvalues for relative, normalized activity, presented in Table 4show greater then unity (from 1.2 to 1.5) values for the‘‘remaining body parts’’. Although for Am and Pu the highestconcentrations were found in internal organs, one cannotexclude the influence of the remains of food (i.e. humus) inthe digestion system, which overestimates the amount of (forinstance) Pu or Am in insect bodies. However, the ‘‘remainingbody parts’’ also contain some inner organs and no digestionsystem, so it is seen that not only radiocaesium or radio-strontium, but also plutonium and americium are incorporatedinto bodies of insects. The 90Sr is shared by ‘‘legs’’, ‘‘innerorgans’’ and ‘‘remaining body parts’’. Caesium and potassiumisotopes follow a similar pattern to each other, with lowconcentrations in legs and heads.

Conclusions

Radionuclides, even those usually considered as refractory,such as plutonium, americium and curium were foundincorporated into inner organs of examined insects. Thedistribution pattern for Pu and Am within the insect bodywas different from that of 90Sr. Both 137Cs and 40K show thirddistribution patterns. Pu and Am seem to cumulate mainly ininternal organs and other body parts. 90Sr is present in internalorgans, body parts, legs, but not in the head, whereas 137Cs and40K were found more evenly distributed with minima for legsand head.The activities of radionuclides connected with hot particle

fractions of Chernobyl fallout were found to be significantlyhigher for samples collected in north-eastern Poland than thosefor samples from southern Poland. The samples analyzedretrospectively, collected over 10 years ago (a few years afterthe Chernobyl accident) combined with recently collectedsamples, showed only a very slight increasing trend of Pu orAm activity, which suggests that even 10 years ago, Pu or Amfrom hot particles were already similarly bio-available as theyare now.The conducted investigation suggests that the beetle species

Anoplotrupes stercorosus is suitable for bio-monitoring ofradioactive contamination. However, appropriate transferfactors should be a matter of concern.

Acknowledgements

The authors are grateful to the Polish State Committee forScientific Research for partial support of this investigation,grant no. 6P04G 07520.

References

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4 P. Szwałko, PhD Thesis, Agricultural University of Krakow, 2001,pp. 198 1 112.

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Table 3 Percentage of activity observed in particular parts of adult beetle body of Anoplotrupes stercorosus

Body part 137Cs (%) 40K (%) 2391240Pu (%) 90Sr (%) 241Am (%)

Heads 9.0 ¡ 0.6 7.9 ¡ 0.8 3.2 ¡ 0.9 1.1 ¡ 0.4 1.9 ¡ 2.9Elytra 9.7 ¡ 0.9 9.3 ¡ 1.0 3.1 ¡ 0.9 3.5 ¡ 5.4 6.9 ¡ 4.7Legs 17.8 ¡ 1.2 17.0 ¡ 1.8 1.5 ¡ 0.7 35.5 ¡ 9.4 1.0 ¡ 0.3Internal organs of thorax and abdomen 19.9 ¡ 1.3 16.7 ¡ 1.7 44.0 ¡ 5.2 19.5 ¡ 4.5 45.6 ¡ 7.1Remaining body parts 43.7 ¡ 1.7 49.1 ¡ 3.2 48.2 ¡ 5.8 40.3 ¡ 5.0 44.7 ¡ 11.5

Table 4 Ratio of percentage of activity observed in particular parts of Anoplotrupes stercorosus body normalized to the percentage of the organ/tissue to whole body (ashed) mass

Body part m (%) 137Cs 40K 2391240Pu 90Sr 241Am

Heads 10.3 0.87 ¡ 0.06 0.77 ¡ 0.08 0.32 ¡ 0.09 0.11 ¡ 0.04 0.18 ¡ 0.28Elytra 8.7 1.11 ¡ 0.10 1.07 ¡ 0.11 0.36 ¡ 0.10 0.41 ¡ 0.62 0.79 ¡ 0.54Legs 21.9 0.81 ¡ 0.06 0.77 ¡ 0.08 0.07 ¡ 0.03 1.62 ¡ 0.43 0.04 ¡ 0.01Internal organs of thorax and abdomen 12.7 1.56 ¡ 0.10 1.31 ¡ 0.14 3.45 ¡ 0.41 1.53 ¡ 0.36 3.58 ¡ 0.55Remaining body parts 32.5 1.34 ¡ 0.05 1.51 ¡ 0.10 1.48 ¡ 0.18 1.24 ¡ 0.15 1.37 ¡ 0.35

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