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Life in a Contaminated Environment: How Soil Nematodes Can Indicate Long-Term Heavy-Metal Pollution

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Figure 1

Map of selected localities: (A) control meadows situated 3.6 km before pollution source; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.
Map of selected localities: (A) control meadows situated 3.6 km before pollution source; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.

Figure 2

RDA biplot of prevailing nematode genera and heavy metals contents in the five alluvial meadows along the Litavka River in the vicinity of the Příbram mine pollution sources. (A) a control meadow 3.6 km upstream of the source of pollution; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source. Amplimer = Amplimerlinius; Acrobelo = Acrobeloides; Aphelench = Aphelenchoides; Aporcela = Aporcelaimellus; Crassolb = Crassolabium; Criconem = Criconemoides; Dipttherp = Diptherophora; Eucephal = Eucephalobus; Eudoryla = Eudorylaimus; Eumonhys = Eumonhystera; Helicotl = Helicotylenchus; Hemicycl = Hemicycliophora; Heterocp = Heterocephalobus; Mesodorl = Mesodorylaimus; Mesorhab = Mesorhabditis; Paratyln = Paratylenchus; Pratylech = Pratylenchoides; Prismatl = Primatolaimus; Teratocep = Teratocepahlobus; Trichodr = Trichodorus; Tylenchr = Tylenchorhynchus. RDA, redundancy analysis.
RDA biplot of prevailing nematode genera and heavy metals contents in the five alluvial meadows along the Litavka River in the vicinity of the Příbram mine pollution sources. (A) a control meadow 3.6 km upstream of the source of pollution; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source. Amplimer = Amplimerlinius; Acrobelo = Acrobeloides; Aphelench = Aphelenchoides; Aporcela = Aporcelaimellus; Crassolb = Crassolabium; Criconem = Criconemoides; Dipttherp = Diptherophora; Eucephal = Eucephalobus; Eudoryla = Eudorylaimus; Eumonhys = Eumonhystera; Helicotl = Helicotylenchus; Hemicycl = Hemicycliophora; Heterocp = Heterocephalobus; Mesodorl = Mesodorylaimus; Mesorhab = Mesorhabditis; Paratyln = Paratylenchus; Pratylech = Pratylenchoides; Prismatl = Primatolaimus; Teratocep = Teratocepahlobus; Trichodr = Trichodorus; Tylenchr = Tylenchorhynchus. RDA, redundancy analysis.

Figure 3

Plots of enrichment vs. structure indices associated with investigated alluvial meadows along the Litavka River contaminated by heavy metals in the vicinity of the Příbram mine pollution sources. A – control; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.
Plots of enrichment vs. structure indices associated with investigated alluvial meadows along the Litavka River contaminated by heavy metals in the vicinity of the Příbram mine pollution sources. A – control; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.

Figure 4

Functional metabolic footprints of nematode associated with investigated alluvial meadows along the Litavka River contaminated by heavy metals in the vicinity of the Příbram mine pollution sources. The vertical axis of each footprint represents the enrichment footprint and the horizontal axis represents the structural footprint. The x-axis coordinates of the metabolic footprint are calculated as SI – 0.5Fs/k and SI + 0.5Fs/k, where Fs is the sum of standardized C utilization by structure indicator taxa. The y-axis coordinates are calculated as EI – 0.5Fe/k and EI + 0.5 Fe/k, where Fe is the sum of standardized C utilization by enrichment indicator taxa. The functional metabolic footprint is depicted by sequentially joining points: SI – 0.5 Fs/k, EI; SI, EI + 0.5 Fe/k; SI + 0.5 Fs/k, EI; SI, EI – 0.5 Fe/k; and SI – 0.5 Fs/k, EI. A – control; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.
Functional metabolic footprints of nematode associated with investigated alluvial meadows along the Litavka River contaminated by heavy metals in the vicinity of the Příbram mine pollution sources. The vertical axis of each footprint represents the enrichment footprint and the horizontal axis represents the structural footprint. The x-axis coordinates of the metabolic footprint are calculated as SI – 0.5Fs/k and SI + 0.5Fs/k, where Fs is the sum of standardized C utilization by structure indicator taxa. The y-axis coordinates are calculated as EI – 0.5Fe/k and EI + 0.5 Fe/k, where Fe is the sum of standardized C utilization by enrichment indicator taxa. The functional metabolic footprint is depicted by sequentially joining points: SI – 0.5 Fs/k, EI; SI, EI + 0.5 Fe/k; SI + 0.5 Fs/k, EI; SI, EI – 0.5 Fe/k; and SI – 0.5 Fs/k, EI. A – control; (B) meadow near waste sedimentation pond; (C) meadow near the heap of mine waste situated 680 m from (B); and (D) meadows near old bakery mill situated 3.7 km from (B) and (E) meadow close to village Jince situated 13.3 km from the pollution source.

Spearman’s rank correlation coefficients between soil physico-chemical properties, heavy metals, and nematode parameters in the alluvial meadows in the vicinity of the Příbram mine along the Litavka River.

As Cd Cr Cu Ni Pb Zn pH/KCl Ntot Cox SM
Abundance Ns ns ns ns ns −0.862** −0.748** −0.659** ns ns ns
No. genera ns ns ns ns ns ns −0.451* ns ns ns ns
Ba1 −0.524* ns ns ns ns −0.599* −0.482* ns ns ns ns
Ba2 −0.633** ns ns ns ns −0.826** −0.859** 0.467* 0.561* ns ns
Ba3 −0.489* ns ns ns 0.442* −0.631** −0.745** ns ns ns 0.412*
Ba4 ns ns ns ns ns −0.455* ns ns ns ns ns
P3 −0.583* ns −0.409* −0.460* ns −0.741** −0.692** ns ns ns ns
P4 ns ns ns ns ns ns −0.423* ns ns ns ns
P5 ns ns ns −0.559* ns −0.699** −0.711** ns ns ns 0.431*
Fu2 −0.507* ns ns ns ns −0.853** −0.626** −0.492* ns ns ns
Fu3 ns ns ns ns 0.455* ns ns −0.522* ns ns ns
Fu4 −0.625** ns ns −0.474* ns −0.769** −0.834** −0.439* ns ns ns
Om3 ns ns ns ns ns ns ns ns ns ns ns
Om4 −0.569* ns ns ns ns −0.845** −0.869** ns −0.422* ns ns
Om5 −0.725** ns ns ns ns −0.766** −0.683** −0.426* −0.445* ns ns
Pp2 ns ns ns ns ns −0.432* −0.457* ns ns ns ns
Pp3 ns ns ns 0.466* 0.529* ns 0.513* ns ns ns 0.436*
Pp4 ns ns ns ns ns ns ns ns ns ns ns
Pp5 ns ns ns ns ns ns ns ns ns ns ns
H´spp ns ns ns −0.425* ns −0.722** −0.634** ns ns ns ns
MI ns ns ns ns ns −0.584** −0.605** 0.433* ns ns ns
MI (2–5) ns ns ns ns ns ns −0.552* ns ns ns ns
PPI ns ns ns ns ns ns ns ns ns ns ns
CI ns ns ns ns ns ns ns ns ns ns ns
EI ns ns ns ns ns −0.439* ns ns ns ns ns
SI −0.548* ns ns ns ns −0.658** −0.742** ns ns ns ns

Mean (±SD) abundance of nematode trophic groups and functional guilds associated with alluvial meadows in the vicinity of the Příbram mine along the Litavka River.

A B C D E
Bacterivores 1160.6 ± 604.5a 218.6 ± 56.3b 115.6 ± 44.9b 139.9 ± 56.8b 284.3 ± 119.9b
Ba1 466.1 ± 142.5a 48.5 ± 31.2b 28.4 ± 18.3b 14.8 ± 10.2b 57.6 ± 30.0b
Ba2 623.3 ± 149.5a 152.3 ± 41.3b 61.9 ± 8.3b 109.3 ± 54.1b 182.3 ± 78.9b
Ba3 62.1 ± 60.6a 17.1 ± 13.8b 19.1 ± 15.3b 15.7 ± 25.1b 32.3 ± 27.2ab
Ba4 9.1 ± 18.1a 0.6 ± 1.1b 0.6 ± 1.1b - 10.0 ± 11.7a
Predators 87.6 ± 79.3 8.8 ± 6.8 12.4 ± 9.8 10.1 ± 6.1 37.1 ± 14.8
P3 7.7 ± 9.2a 2.9 ± 5.8a 2.9 ± 5.8a 1.9 ± 3.8a -
P4 19.5 ± 12.1a 5.1 ± 5.0b 12.4 ± 9.8a 8.1 ± 5.0a 33.3 ± 10.8a
P5 60.4 ± 40.2a 0.8 ± 1.6b - - 3.8 ± 4.5b
Fungivores 276.1 ± 196.9a 49.0 ± 26.7b 25.3 ± 7.6b 30.6 ± 10.9b 60.6 ± 24.8b
Fu2 150.3 ± 39.8a 39.4 ± 15.1b 16.8 ± 7.9b 24.3 ± 7.1b 46.6 ± 31.4b
Fu3 10.9 ± 14.9a 5.0 ± 5.0a 8.5 ± 7.9a 6.2 ± 6.5a 6.0 ± 8.2a
Fu4 114.8 ± 46.3a 4.6 ± 8.0b 4.6 ± 8.0b - 7.6 ± 10.2b
Omnivores 220.7 ± 167.4a 17.2 ± 6.6b 7.9 ± 3.0b 18.1 ± 19.0b 85.0 ± 44.3ab
Om3 3.0 ± 6.1 - - - -
Om4 203.6 ± 90.0a 12.7 ± 10.1b 5.9 ± 3.7b 14.3 ± 14.7b 63.9 ± 40.6b
Om5 14.0 ± 6.4a 4.3 ± 5.0b 2.0 ± 2.4b 3.8 ± 4.4b 21.1 ± 5.6a
Plant parasites 451.8 ± 117.5a 299.7 ± 66.9a 351.4 ± 98.0a 782.5 ± 260.8a 893.8 ± 249.3a
Pp2 108.3 ± 52.5a 84.6 ± 40.5a 44.4 ± 25.7b 187.3 ± 121.4a 397.7 ± 262.4a
Pp3 340.1 ± 15.7a 206.1 ± 67.4a 308.0 ± 84.9a 589.2 ± 206.3a 540.5 ± 111.2a
Pp4 - 8.4 ± 6.7 - - 45.5 ± 35.8
Pp5 - 3.4 ± 4.3 - - 23.8 ± 28.9

Soil physico- chemical properties (mean ± SD) associated with alluvial meadows in the vicinity of the Příbram mine along the Litavka River.

Parameter Sites
A B C D E
pH/KCl 6.89 ± 0.25a 5.42 ± 0.21a 5.20 ± −0.11b 5.56 ± 0.16b 6.75 ± 0.18a
Soil moisture (%) 35.2 ± 5.6a 31.5 ± 2.9a 29.5 ± 3.1a 30.5 ± 3.3a 31.5 ± 2.8a
Ntot (%) 0.87 ± 0.15a 0.61 ± 0.10b 0.74 ± 0.32b 0.66 ± 0.15b 0.83 ± 0.22a
Cox (%) 14.25 ± 1.69a 8.25 ± 2.19b 10.67 ± 3.61b 12.29 ± 2.69a 12.87 ± 3.16a

Total concentrations of heavy metals (mg/kg) (mean ± SD) associated with alluvial meadows in the vicinity of the Příbram mine along the Litavka River.

Element Sites
Max
A B C D E
As 38.2 ± 3.3a 688.8 ± 91.8c 677.4 ± 181.1c 176.2 ± 47.7b 31.8 ± 17.8a 20
Cd 1.9 ± 0.2a 19.5 ± 1.9b 51.5 ± 23.1c 28.4 ± 12.9bc 2.1 ± 1.0a 0.5
Cr 37.9 ± 5.1a 58.2 ± 1.3b 68.8 ± 59.9a 40.4 ± 5.7a 28.7 ± 1.7a 90
Cu 35.7 ± 5.3a 97.6 ± 8.7b 139.6 ± 24.6c 61.9 ± 30.7ab 23.1 ± 11.9a 60
Ni 19.1 ± 1.3a 38.1 ± 2.4b 33.2 ± 8.2ab 22.9 ± 8.3a 15.4 ± 1.8a 50
Pb 553.3 ± 237.6a 3734.1 ± 1915.6b 4030.9 ± 165.1bc 2682.2 ± 460.3b 689.8 ± 474.3a 60
Zn 346.2 ± 34.6a 2949.5 ± 435.9b 6867.8 ± 1374.9c 3866.4 ± 1146.2b 566.4 ± 490.1a 120

Total nematode abundance, number of genera, nematode community indices associated with alluvial meadows in the vicinity of the Příbram mine along the Litavka River (mean ± SD).

Indices A B C D E
Nematode abundance 2193.2 ± 358.9a 595.8 ± 130.9b 513.8 ± 122.3b 974.9 ± 340.4b 1674.1 ± 263.5a
Genera number 31.8 ± 1.7a 25.8 ± 1.5b 27.5 ± 5.1a 21.5 ± 3.1b 30.5 ± 4.2a
Maturity Index 2.68 ± 0.12a 2.12 ± 0.09b 2.20 ± 0.10b 2.22 ± 0.21b 2.63 ± 0.15a
Maturity Index (2-5) 2.89 ± 0.07a 2.33 ± 0.16b 2.37 ± 0.15b 2.33 ± 0.19b 2.85 ± 0.22a
Plant Parasitic Index 2.69 ± 0.11a 2.80 ± 0.18a 2.88 ± 0.04a 2.77 ± 0.14a 2.73 ± 0.16a
Diversity Index (H´gen) 3.49 ± 0.15a 2.13 ± 0.25b 2.47 ± 0.26b 2.22 ± 0.18b 3.09 ± 0.10a
Channel Index 8.8 ± 4.0a 23.0 ± 11.8b 36.4 ± 12.9b 30.9 ± 6.6b 22.9 ± 19.6ab
Basal Index 16.6 ± 2.3a 32.9 ± 8.3b 33.9 ± 10.7b 33.7 ± 4.6b 19.9 ± 9.3a
Enrichment Index 71.5 ± 4.4a 53.2 ± 10.9b 50.3 ± 11.5b 37.9 ± 16.2b 66.7 ± 10.1b
Structure Index 72.0 ± 3.2a 44.5 ± 18.7b 49.8 ± 13.1b 47.9 ± 16.2b 73.8 ± 12.3a
Total biomass, mg 7.5 ± 1.2a 2.1 ± 1.2b 1.9 ± 1.3b 4.9 ± 2.9b 8.6 ± 2.5a

A abundance of nematode genera (mean ± SD) associated with alluvial meadows in the vicinity of the Příbram mine along the Litavka River.

Genus/trophic group c-p A B C D E
Bacterivores
Mesorhabditis 1 251.6 ± 103.4 24.3 ± 13.0 22.6 ± 14.2 5.3 ± 8.3 24.8 ± 14.5
Plectus 2 241.3 ± 44.1 23.9 ± 20.7 10.5 ± 2.9 0.9 ± 0.5 15.4 ± 2.8
Rhabditis 1 187.0 ± 63.9 21.8 ± 17.7 5.9 ± 1.2 9.5 ± 11.6 26.3 ± 10.8
Acrobeloides 2 143.3 ± 75.1 72.1 ± 8.4 15.4 ± 7.9 51.1 ± 26.6 40.5 ± 35.6
Cephalobus 2 123.5 ± 61.0 12.1 ± 8.9 5.9 ± 3.9 9.6 ± 10.2 23.2 ± 18.7
Eucephalobus 2 53.3 ± 27.8 12.2 ± 9.3 20.5 ± 10.8 22.5 ± 15.9 11.6 ± 10.1
Prismatolaimus 3 51.1 ± 60.8 12.1 ± 7.9 7.0 ± 3.3 11.9 ± 17.5 17.6 ± 5.3
Bunonema 1 27.4 ± 14.9 - - - 2.2 ± 4.4
Eumonhystera 2 26.2 ± 24.1 4.6 ± 5.9 5.0 ± 2.4 - 4.4 ± 8.7
Heterocephalobus 2 18.3 ± 5.6 4.5 ± 3.4 - 8.6 ± 3.9 -
Alaimus 4 9.1 ± 18.1 0.6 ± 1.1 4.3 ± 7.3 - 12.2 ± 9.8
Bastiania 3 8.7 ± 5.9 2.1 ± 4.2 2.6 ± 1.9 1.9 ± 3.8 4.6 ± 9.3
Anaplectus 2 8.0 ± 5.4 1.9 ± 1.4 - - -
Cervidellus 2 7.9 ± 15.8 - - 8.1 ± 6.6 47.0 ± 12.8
Teratocephalus 3 2.3 ± 4.6 1.4 ± 2.9 9.5 ± 2.6 1.9 ± 3.8 10.3 ± 5.9
Eudiplogaster 2 1.5 ± 3.0 0.5 ± 1.0 - 1.9 ± 3.8 2.3 ± 4.6
Acrobeles 2 - - - 1.9 ± 3.8 16.8 ± 11.5
Amphidelus 4 - - 1.9 ± 3.8 - -
Ceratoplectus 2 - 1.1 ± 2.3 2.2 ± 2.5 - -
Diploscapter 1 - 1.4 ± 1.7 - - 2.2 ± 4.4
Euteratocephalus 2 - 4.0 ± 3.8 3.0 ± 2.0 - -
Chiloplacus 2 - - - 2.9 ± 3.7 8.6 ± 10.6
Panagrolaimus 1 - 1.0 ± 2.1 - - 2.2 ± 4.4
Prodesmodora 3 - 1.5 ± 2.0 - - -
Wilsonema 2 - 15.4 ± 14.8 - 1.9 ± 2.1 12.4 ± 8.3
Fungivores
Dorylaimoides 4 104.9 ± 59.4 - - - -
Aphelenchoides 2 113.6 ± 54.2 7.9 ± 5.9 8.2 ± 6.6 9.5 ± 11.5 39.6 ± 7.8
Aphelenchus 2 24.4 ± 37.3 20.9 ± 12.3 2.1 ± 3.0 11.9 ± 6.2 2.2 ± 4.4
Ditylenchus 2 12.5 ± 15.4 9.3 ± 7.6 6.5 ± 3.0 2.9 ± 5.9 3.2 ± 6.5
Diptheropthora 3 10.9 ± 14.9 5.0 ± 5.0 8.5 ± 4.5 6.2 ± 2.8 6.0 ± 4.2
Tylencholaimus 4 5.4 ± 6.3 4.2 ± 8.3 - - 7.6 ± 10.2
Tylencholaimellus 4 4.5 ± 9.1 0.5 ± 1.0 - - -
Paraphelenchus 2 - 1.2 ± 2.5 - - 1.6 ± 3.2
Omnivores
Eudorylaimus 4 91.1 ± 61.5 3.0 ± 3.9 0.9 ± 0.9 2.9 ± 3.7 53.1 ± 38.5
Mesodorylaimus 4 76.2 ± 26.6 - - - 4.4 ± 8.7
Oxydirus 5 60.5 ± 40.2 0.8 ± 1.6 - - 1.6 ± 3.2
Dorylaimus 4 24.3 ± 40.7 0.5 ± 1.0 2.4 ± 1.8 - 1.6 ± 3.2
Aporcelaimellus 5 13.9 ± 16.4 1.0 ± 2.1 2.0 ± 1.4 3.8 ± 4.1 21.1 ± 6.9
Pungentus 4 10.7 ± 12.6 0.4 ± 0.8 0.5 ± 1.0 - 4.9 ± 5.7
Achromadora 3 3.1 ± 6.1 - - - -
Crassolabium 4 1.5 ± 3.0 6.7 ± 6.3 1.5 ± 1.9 1.9 ± 1.2 -
Axonchium 5 - 2.8 ± 5.7 - - -
Discolaimium 4 - 2.1 ± 4.2 - - -
Microdorylaimus 4 - - 0.5 ± 1.1 9.5 ± 11.4 -
Paractinolaimus 5 - 0.4 ± 0.8 - - -
Predators
Tripyla 3 7.7 ± 9.1 2.9 ± 5.9 - 1.9 ± 3.8 -
Mononchus 4 7.3 ± 9.0 0.5 ± 1.0 3.4 ± 2.1 - -
Prionchulus 4 6.2 ± 8.7 - - - 1.6 ± 2.1
Clarkus 4 3.1 ± 6.1 0.9 ± 1.0 8.0 ± 4.6 8.1 ± 5.0 14.4 ± 9.8
Mylonchulus 4 3.1 ± 6.2 3.7 ± 5.9 1.1 ± 2.1 - 17.3 ± 14.6
Nygolaimus 5 - - - - 2.2 ± 4.4
Plant parasites
Helicotylenchus 3 163.8 ± 82.9 38.7 ± 31.1 60.9 ± 21.6 85.7 ± 64.8 118.0 ± 57.6
Heterodera 3 57.9 ± 47.4 31.8 ± 2.4 27.8 ± 17.3 78.6 ± 21.8 52.3 ± 134.8
Pratylenchoides 3 36.9 ± 30.1 5.0 ± 1.2 - - -
Malenchus 2 50.9 ± 52.7 9.2 ± 2.5 5.8 ± 2.4 95.5 ± 21.9 2.2 ± 4.4
Rotylenchus 3 29.3 ± 31.7 - 0.5 ± 1.1 0.9 ± 1.9 -
Geocenamus 3 27.0 ± 20.3 67.7 ± 47.0 62.5 ± 16.3 53.8 ± 4.4 22.4 ± 16.6
Gracilacus 2 26.6 ± 12.9 39.8 ± 33.0 - - 7.0 ± 13.9
Filenchus 2 15.3 ± 12.0 5.1 ± 4.7 0.9 ± 1.9 4.3 ± 3.3 9.2 ± 10.6
Tylenchorhynchus 3 14.7 ± 12.0 23.9 ± 27.1 3.9 ± 5.1 - 4.9 ± 5.7
Meloidogyne 3 7.5 ± 8.2 10.7 ± 26.2 - 20.5 ± 14.5 99.6 ± 33.9
Tylenchus 2 6.8 ± 9.9 0.8 ± 1.6 0.5 ± 1.1 1.9 ± 3.8 -
Coslenchus 2 6.1 ± 12.2 2.8 ± 3.7 - - 2.3 ± 4.6
Amplimerlinius 3 3.0 ± 6.1 1.3 ± 2.0 - - -
Lelenchus 2 2.6 ± 5.3 - - 3.8 ± 7.6 -
Criconemoides 3 - - - - 9.7 ± 7.0
Hemicycliophora 3 - - 105.7 ± 49.2 303.1 ± 63.3 195.3 ± 110.3
Longidorus 5 - 3.4 ± 4.3 - - 23.8 ± 8.2
Mesocriconema 3 - - 1.6 ± 2.1 - 15.6 ± 12.1
Paratrichodorus 4 - 4.1 ± 1.5 - - 24.9 ± 30.2
Paratylenchus 2 - 27.0 ± 16.2 33.3 ± 10.8 23.4 ± 9.7 177.1 ± 159.5
Pratylenchus 3 - 31.6 ± 8.4 45.1 ± 14.1 107.1 ± 28.9 122.8 ± 40.2
Trichodorus 4 - 4.3 ± 5.9 - - 20.5 ± 18.4
Xenocriconemella 3 - 4.3 ± 7.2 - - -
Aglenchus 2 - - - 3.9 ± 2.5 -
Boleodorus 2 - - 2.8 ± 3.5 - -
Psilenchus 2 - - 1.1 ± 1.2 - -
Insect parasites
Steinernema 1 - - 2.8 ± 5.7 3.8 ± 4.4
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