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Table 1. Predatory mites collected during the survey
Sl. No. Predatory mite Order Family
1 . Neoseiulus longispinosus Evans Acari Phytoseiidae
2 . Neoseiulus herbicolus Chant Acari Phytoseiidae
3 . Neoseiulus rhabdus Denmark Acari Phytoseiidae
4 . Bdella sp. Acari Bdellidae
5. Agistemus sp. Acari Stigmaeidae
6. Tydeius sp. Acari Tydeidae
Table 2. Developmental time (in days) (Mean±SE) of immature stages of N. longispinosus at five constant temperaturesa
Temperature (ºC)
nb Sex Egg Larva Protonymph Deutonymph Total Intersexual differencec
(P) 15 14
18 Male
Female 3.7±0.03a 3.8±0.05a
2.9±0.04a 2.9±0.03a
3.0±0.03a 3.1±0.04a
4.3±0.05a 4.3±0.05a
13.9±0.11a 14.0±0.07a
0.467
20 12 17
Male Female
2.4±0.07b 2.5±0.05b
0.9±0.04b 0.9±0.03b
2.9±0.05b 2.9±0.04b
3.1±0.03b 3.1±0.06b
9.3±0.10b 9.4±0.08b
0.273
25 11 22
Male Female
1.8±0.08c 1.8±0.04c
0.6±0.02c 0.6±0.02c
1.3±0.03c 1.3±0.03c
1.3±0.04c 1.3±0.02c
4.9±0.08c 4.9±0.07c
0.347
30 13 19
Male Female
1.3±0.05d 1.4±0.03d
0.5±0.02c 0.5±0.01c
1.0±0.03d 1.0±0.03d
1.2±0.03c 1.3±0.03c
4.0±0.07d 4.2±0.05d
0.036
35 47 Not known
1.1±0.16e Didn’t survive
- - - -
aMeans with in the same column followed by same letters are not significantly different at P<0.05, Student-Newman-Keuls test. b n= number of samples cIntersexual difference in total developmental duration was analyzed using Mann-Whitney U-test
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Table 3. Hatchability and survival rate of immature stages of N. longispinosus at five constant temperatures
Temperature (ºC)
No. of eggs tested
Hatchability (%) No. of larvae introduced
Maturity (%) Mortality (egg-to-adult)
(%) 15 100 98 98 96.94 5.00 20 100 99 99 96.96 4.00 25 100 98 98 96.94 5.00 30 100 97 97 96.91 6.00 35 100 73 73 0 100
Table 4. Linear regression analysis of developmental rate and temperature, and estimated lower developmental threshold and thermal constant for N. longispinosus
Sex/Stage Regression equationa
r2 Lower development
threshold (t0, ºC) (±SE)b
Thermal constant (K)
(degree-days) (±SE)b
Female Egg y= - 0.1926+0.0300x 0.9990 06.4±0.1 33.3±0.7 Larva y= - 1.0686+0.1036x 0.9205 10.3±0.9 09.7±2.0 Protonymph y= - 0.4852+0.0487x 0.9112 10.0±0.9 20.5±4.5 Deutonymph y= - 0.4386+0.0432x 0.8734 10.2±1.1 23.2±6.2 Egg-Adult y= - 0.1140+0.0119x 0.9605 10.0±0.7 84.0±12.0 Male Egg y= - 0.2390+ 0.0331x 0.9942 07.2±0.2 30.2±1.6 Larva y= - 1.1567+ 0.1093x 0.9205 10.6±0.9 09.2±1.9 Protonymph y= - 0.5099+ 0.0503x 0.9092 10.1±1.0 19.9±4.4 Deutonymph y= - 0.4458+ 0.0436x 0.8879 10.2±1.1 23.0±5.8 Egg-Adult y= - 0.1234+ 0.0125x 0.9655 09.9±0.6 80.0±10.7 aSimple linear regression analysis was applied to the developmental data within 15-30ºC bStandard errors were calculated according to the equations given by Campbell et al. (1974).
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Table 5. Duration (in days) and rate of reproduction (Mean±SE) of N. longispinosus adult females at three constant temperaturesa
Parameter Temperature (ºC)
20(n=12) 25(n=21) 30(n=11) (χ2-value)b Pre-oviposition period 2.0±0.2a 1.2±0.1b 1.0±0.0b 18.31 Oviposition period 42.1±0.7a 36.1±0.7b 23.4±0.8c 34.47 Post oviposition period 6.1±0.4a 2.1±0.2b 1.9±0.2b 26.96 Total No. of eggs/female 71.8±0.9a 62.1±0.7b 60.9±1.1b 26.06 Average No. of eggs/female /day 1.7±0.03b 1.7±0.1b 2.6±0.1a 24.30 Mean male longevity 55.6±0.9a 44.9±0.5b 30.9±1.0c 36.72 Mean female longevity 50.9±0.4a 39.1±0.6b 25.9±0.6c 36.93 Sex ratio (Female: Male) 1:0.50 1:0.53 1:0.47 aMeans with in a row followed by same letters are not significantly different at P<0.05, Student-Newman-Keuls test). n=number of samples b Kruskal-wallis test, P<0.001
Table 6. Life table parameters of N. longispinosus on O. coffeae
Parameter 20 ºC 25 ºC 30 ºC Intrinsic rate of natural increase (rm/day) 0.132 0.219 0.268 Net reproductive rate (R0) 40.7 35.9 35.5 Gross reproductive rate (Σmx) 47.9 40.5 41.4 Mean generation time (T) 28.2 16.4 13.3 Finite rate of increase (λ) 1.1 1.2 1.3 Weekly multiplication 2.5 4.6 6.5 Doubling time (DT) 5.3 3.2 3.2
Table 7. Prey consumption of N. longispinosus on O. coffeae
Predator stages Mean No. of red spider mite consumed±SE* (n=10)
Egg Larva Protonymph Deutonymph Adult Larva 3.8±0.2a 9±0.3a 5.5±0.3a 0a 0a Protonymph 5.1±0.3b 16.2±0.7bc 6.2±0.3a 9.0±0.6b 3.1±0.3c Deutonymph 6.1±0.5b 17.2±0.6c 12.0±0.6b 11.2±0.4c 4.0±0.4d Adult male 4.0±0.4a 14.8±0.7b 12.1±0.6b 9.9±0.6bc 1.9±0.3b Adult female 13.3±0.5c 21.9±0.6d 18.3±0.5c 15.9±0.7d 5.5±0.3e * Prey consumption of larvae was recorded 10 h after the commencement of the experiment and that of other stages after 24 h. The means followed by same letter in the same column are not significantly different at 0.05 level as determined by Duncan’s Multiple Range Test.
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Table 8. Functional response of N. longispinosus adult females to O. coffeae adults at different temperatures
Parameters* 10 ºC 15 ºC 20 ºC 25 ºC 30 ºC 35 ºC
Na 0.563 N 1 + 0.174 N
0.687 N 1 +0.196 N
0.991 N 1 + 0.277 N
1.230 N 1 + 0.266 N
1.187 N 1 +0.204 N
1.183 N 1 + 0.202 N
Th 0.309 0.285 0.280 0.216 0.172 0.171 a 0.563 0.687 0.991 1.230 1.187 1.183 a/Th 1.819 2.415 3.546 5.685 6.892 6.912 K 3.232 3.514 3.578 4.623 5.807 5.845 * Na = number of prey consumed; N = initial prey density; Th = handling time; a = attack rate; K = maximum predation rate
Table 9. Functional response of N. longispinosus adult females to different stages of O. coffeae Parameters* Egg Larva Protonymph Deutonymph Adult female
Na 0.826 N
1 + 0.044 N 1.012 N
1 + 0.015 N 1.022 N
1 + 0.036 N 1.032 N
1 + 0.047 N 1.211 N
1 + 0.237 N Th 0.053 0.015 0.035 0.046 0.196 a 0.826 1.012 1.022 1.032 1.211 a/Th 15.59 67.47 28.878 22.43 6.181 K 18.87 66.67 28.25 21.74 5.102 * Na = number of prey consumed; N = initial prey density; Th = handling time; a = attack rate; K = maximum predation rate
Table 10. Numbers of predator and prey remaining after 5 days on the leaf lets received various ratios of predator: prey in laboratory
No. of predators Predator: Prey ratio Number of predator Mean* ± SE
Number of prey Mean* ± SE
1 1:100 8.4±0.4d 136.8±5.7b 2 1:50 16.4±1.4ab 84.2±3.0bc 3 1:33 17.8±1.2a 49.2±1.6c 4 1:25 14.4±0.5b 20.0±1.1c 5 1:20 15.4±0.8b 19.8±1.4c 10 1:10 11.2±0.4c 9.0±0.7c 20 1:5 14.6±0.5b 1.2±0.4c 0 Control 0e 335.0±75.1a
*Mans followed by the same letter in the same column are not significantly different at 0.05 level as determined by DMRT
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Table 11. Numbers of predator and prey remaining after 20 days on the potted plant received various ratios of predator: prey in green house
No. of predators Predator: Prey ratio Number of predator (Mean* ± SE)
Number of prey (Mean* ± SE)
2 1:100 3.0±0.6cd 477.0±11.3b 4 1:50 6.0±2.1bc 264.7±28.76c 8 1:25 9.3±2.0ab 109.3±3.8d 20 1:10 14.3±2.7a 61.7±8.1e 0 Control 0d 593.0±7.6a
*Means followed by the same letter in the same column are not significantly different at 0.05 level as determined by DMRT
Table 13. Volatile compounds emitted from O. coffeae infested leaves S. No. Retention time (RT) Name of the compound from library Molecular weight
1. 8.05 Undecane 156 2. 11.94 Alpha pinene 136 3. 12.54 1,3,6-Octatriene, 3,7-dimethyl-, (Z) (Beta-
Ocimene) 136
4. 23.99 1,6-Octadien-3-ol, 3,7-dimethyl (Linalool) 154 5. 30.23 3-ethyl Benzaldehyde 134 6. 31.16 à-Farnesene 204 7. 34.62 Para acetyl ethyl benzene 148
Table 12. Field efficacy of N. longispinosus as a biocontrol agent for O. coffeae No. of
sampling & release
No. of RSM (Mean ± SE)*
No. of N. longispinosus (Mean ± SE)*
Reduction of RSM over control (%)
Control Release Control Release 1a 68.0±5.5a 74.7±4.7d 11.7±1.5a 9.3±0.9bc - 2b 75.7±3.2ab 77.3±3.2d 10.3±3.0a 9.7±1.2bc - 3c 87.3±2.3bc 55.7±3.8c 11.0±1.0a 14.3±0.9d 36.25 4 84.3±6.2abc 30.7±2.4b 11.3±1.2a 16.7±1.5d 63.63 5 98.3±3.4cd 18.3±1.5a 13.7±1.5a 11.0±1.2c 81.36 6 109.7±4.1d 14.7±0.9a 14.3±1.5a 6.7±0.3b 86.62 7 114.3±6.4d 10.7±0.9a 17.3±2.3a 3.0±0.6a 90.67
*Mean numbers of RSM or N. longispinosus per 6 leaves; a Pre-treatment sampling & I release; b first sampling & II release ; c second sampling & III release. 25 bushes/plot, 6 leaves sampling Means within the same column followed by same letters are not significantly different at P<0.05, Student-Newman-Keuls test.
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Table 14. Volatile compounds emitted from normal leaves S. No. Retention time (RT) Name of the compound from library Molecular weight
1. 12.20 1-fluorodecane 188 2. 12.48 Alpha pinene 136 3. 17.83 Tetradecane 198 4. 19.73 2,6,10,15-tetramethylheptadecane 298 5. 25.60 Hexadecane 226 6. 27.33 2,3,5,8-tetramethyldecane 199 7. 30.24 3-ethylbenzaldehyde 134 8. 34.31 2-methyltridecane 198 9. 34.64 Paraacetylethylbenzene 148 10. 45.56 Benzoic acid, 4-ethoxy ethyl ester 194
Table 15. Impact of pesticides on N. longispinosus under laboratory condition-Direct toxicity
Pesticide Mortality (Mean ± SE)* (%) 24 h 48 h 72 h 96 h
Propargite 4.0±2.5a 4.0±2.5ab 8.0±3.7a 8.0±3.7a Fenpyroximate 100±0.0c 100±0.0d 100±0.0d 100±0.0d Ethion 20.0±4.5b 36.0±5.1c 42.0±6.6c 52.0±9.7c Hexythiazox 0a 2.0±2.0ab 4.0±4.0a 6.0±4.0a NKAE 0a 10.0±3.16b 10.0±3.16a 10.0±3.16a P. fumososroseius 0a 0a 4.0±2.45a 4.0±2.5a Lime sulphur 18.0±3.7b 30.0±4.5c 30.0±4.5b 32.0±5.8b Control 0a 0a 0a 0a *Means in the same column followed by the same letter are not significantly different at 0.05 level as determined by DMRT
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Table 17. Impact of pesticides on immature stages of N. longispinosus under laboratory condition
Pesticide Percentage survival up to adult (Mean ± SE)*
Propargite 47.4±6.3c Fenpyroximate 1.0±0a Ethion 17.9±5.0b Hexythiazox 7.4±3.7ab NKAE 63.4±2.9d P. fumososroseius 76.1±7.3d Lime sulphur 93.2±5.3e Control 96.0±2.5e *Means in the same column followed by the same letter are not significantly different at 0.05 level as determined by DMRT
Table 18. Impact of pesticides on N. longispinosus under field condition
Mean No. of red spider mite in the experimental blocks±SE
Pre treatment I week II week III week IV week Fenpyroximate 13.7±0.3a 6.0±1.2a 1.3±0.3a 1.7±0.3a 1.3±0.3a Propargite 17.0±0.6ab 16.0±1.5ab 10.0±1.2b 8.7±1.5abc 6.3±1.2ab Ethion 20.0±3.1ab 15.0±2.1ab 12.3±1.7b 10.3±1.2abc 9.7±1.2b Hexythiazox 21.0±1.2ab 18.3±0.9ab 18.0±1.5b 11.0±1.2abc 8.3±1.5ab NKAE 18.7±2.6ab 15.7±2.0ab 10. 7±0.9b 9.3±0.9abc 9.7±1.5b P. fumosoroseius 24.7±0.7b 23.7±3.7b 14.3±0.9b 7.0±0.6ab 7.7±1.5ab Lime sulphur 18.7±4.3ab 15.3±4.5ab 13.7±3.7b 11.7±3.7bc 12.0±2.3b Control 16.7±1.5ab 18.3±3.2ab 18.7±2.9b 16.7±3.5c 22.0±3.1c Means in the same column followed by the same letter are not significantly different at 0.05 level as determined by DMRT
Table 16. Impact of pesticides on N. longispinosus under laboratory condition-Indirect toxicity
Pesticide Mortality (Mean ± SE)* (%) 24 h 48 h 72 h 96 h
Propargite 0a 0a 0a 0a Fenpyroximate 42.0±3.7c 62.0±2.0d 76.0±2.5d 80.0±3.2d Ethion 24.0±7.5b 40.0±8.37c 44.0±6.8c 48.0±8.6c Hexythiazox 0a 0a 0a 0a NKAE 0a 0a 0a 0a P. fumososroseius 0a 0a 0a 0a Lime sulphur 18.0±3.7b 24.0±2.45b 24.0±2.5b 24.0±2.5b Control 0a 0a 0a 0a *Means in the same column followed by the same letter are not significantly different at 0.05 level as determined by DMRT
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Table 19. Selection of N. longispinosus for resistance to fenpyroximate
No. of selection
Dosage (ml/L)
LC50 (ml/L) (95% CL)a Slope ± SE RR*
1st selection 0.21 0.23 (0.05-0.40) 1.65±0.15
2nd selection 0.33 0.27 (0.04-0.49) 1.43±0.14 1.15
3rd selection 0.40 0.40 (0.09-0.87) 0.99±0.14 1.75
4th selection 0.73 0.53 (0.11-2.84) 0.85±0.13 2.29
5th selection 1.05 0.74 (0.38-3.29) 0.98±0.14 3.22
6th selection 1.35 0.96 (0.53-6.40) 1.11±0.14 4.17
*Resistance Ratio (LC50 of resistant strain/LC50 of susceptible strain) Table 20. Monooxygenase activities in adult female N. longispinosus after selection with fenpyroximate
Population Enzyme activity (mOD/min/ mg-1/ protein ± SD)* R/S*
Control 0.209
Fenpyroximate 6 0.859 4.10
*Enzyme activity of resistant strain/Enzyme activity of susceptible strain (fold of resistance developed)
Table 21. Oviposition of resistant and susceptible strains on pesticide treated and untreated surfaces of leaves
No. of eggs laid (Mean ± SE)
Control mites Resistant mites Treated portion 0±0 10.2±1.39
Untreated portion 11±1.14 7.40±1.33
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Fig. 1. Tea growing areas in the world
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Fig. 2. Tea growing areas in India
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Fig. 3. Tea growing areas in southern India
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Fig.4. Spatial distribution of pests in a tea bush
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Organic field Conventional field
Fig. 5. Effect of organic and conventional field practices on seasonal dynamics of N. longispinosus
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Fig. 6. Age specific survival (lx), age-specific fecundity (mx) and lxmx curves of N. longispinosus females at 20, 25 and 30 ºC.
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Fig. 7. Control of O. coffeae population by N. longispinosus as a function of time -laboratory experiment
Fig. 8. Average numbers (±SE) of O. coffeae eggs and larvae as function of time on experimental arena
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Fig. 9. Prey stage preference (mean ± SE) of different life stages of N. longispinosus on developmental stages of O. coffeae
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Fig. 10. Numerical responses (mean±SE) of N. longispinosus female to different density of O. coffeae females
Fig. 11. Numerical responses (mean ± SE) of N. longispinosus female to O. coffeae adults at different temperatures
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Fig. 12. Chromatogram of volatiles captured from O. coffeae infested leaves
Fig. 13. Chromatogram of volatiles captured from normal tea leaves
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Fig. 14. Response of N. longispinosus to volatiles from RSM infested tea leaves in Y tube olfactometer-effect of starvation
Fig. 15. Response of N. longispinosus to volatiles from RSM infested tea leaves in Y tube olfactometer-effect of the level of infestation
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Fig. 16. Response of N. longispinosus to volatiles from RSM infested tea leaves in Y tube olfactometer-effect of previous infestation, Response of predators to HIPVs from infested plants alone and effect of artificial damage on tea leaves
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Plate 1. Red spider mite infested tea field in contrast to the normal one
Normal tea field Red spider mite infested tea field
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Plate 2. Life cycle of the red spider mite, Oligonychus coffeae
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Plate 3. Mass rearing of red spider mites and N. longispinosus by tray culture
method
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Plate 4. Dynamic head space (DHS) volatile extraction unit
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Plate 5. Experimental setup for behavioral response study
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Plate 6. Life cycle of N. longispinosus
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