Growth and Development of Leptocybe invasa as Affected by Environmental Temperature
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摘要:
目的 致瘿性昆虫—桉树枝瘿姬小蜂（Leptocybe invasa）的生长发育、繁殖及分布主要受到温度的影响，而探明温度对桉树枝瘿姬小蜂生长发育的影响，可为该害虫的虫情监测和预测预报提供科学依据。 方法 采用室内恒温接种、饲养的方法，测定了桉树枝瘿姬小蜂在19、22、25、28、31℃下各虫态的发育历期和发育速率，利用直线回归（图测法）、线性回归和 Logistic 回归方程分析了温度与各虫态发育速率的关系，并用直线回归方程求得各虫态发育起点温度和有效积温。 结果 温度对桉树枝瘿姬小蜂卵至蛹各虫态、全世代的发育历期及成虫寿命均有显著影响，在19～31℃范围内，各虫态发育历期均随温度升高而逐渐缩短，发育速率与温度呈显著正相关。图测法直线回归方程更能准确拟合桉树枝瘿姬小蜂各虫态发育速率与温度之间的关系。桉树枝瘿姬小蜂卵、幼虫、蛹、成虫（存活）及全世代的发育起点温度分别为10.78、10.29、14.64、14.73和12.68℃，有效积温分别为86.96、588.24、92.59、104.17和712.34 d·℃，发育最适温度分别为25.68、25.65、24.58、26.42和23.84℃，发育适宜温区为12.21～35.48℃。2017—2019年，该害虫在江西赣南发生世代数理论值为4.27代，与实际观察结果一致。 结论 温度是影响桉树枝瘿姬小蜂生长发育的重要因素，本研究结果可为预测该蜂的发育历期、发生区域、发生期和发生代数提供科学依据和技术支撑，尤其对该蜂成虫发生期预测预报和有效防控具有重要的指导意义。 Abstract:Objective Correlations between the growth, reproduction, and distribution of Leptocybe invasa, an insect that causes galls in plants, and the surrounding temperature were studied to facilitate the monitoring and controlling the spread of the pest wasp. Method Duration and speed of each development state of the wasp were measured by inoculating and feeding the larvae indoor at 19, 22, 25, 28 and 31℃. Graphic, linear, and logistic regression analyses were applied to establish the correlations. Threshold and effective temperatures for the emergences of various stages of the insect were obtained using a linear regression model. Result Temperature was found to significantly affect not only at the stage from egg to pupa, but also the entire development and adult life of the insect. The increasing temperature hastened the insect development process. The graphic linear regression model seemed to more accurately describe their correlation than the other models. In Gannan, Jiangxi, the threshold emergence temperature of the eggs was found to be 10.78℃; that of the larva, 10.29℃; that of the pupa, 14.64℃; the adult survival threshold temperature, 14.73℃; and, the temperature for the development of an entire generation, 12.68℃. The accumulated effective temperature for the eggs was found to be 86.96℃; that for the larva, 588.24℃; that for the pupa, 92.59℃; that for the adult survival, 104.17℃; and, that for the development of an entire generation, 712.34 d·℃. And, the optimum temperature for the egg development, 25.68℃; for the larvae, 25.65℃; for the pupa, 24.58℃; for the adult survival, 26.42℃; and, for the development of an entire generation, 23.84℃ with a range between 12.21℃ and 35.48℃. The model estimated 4.27 generations of L. invasa had taken place between 2017 and 2019 which was in agreement with what was observed in the field. Conclusion Temperature was critical for the development of L. invasa. Understanding the correlation would aid in predicting the development, infestation time and area, and emerging generations of L. invasa for effective control of the pest. -
图 1 桉树枝瘿姬小蜂各虫态发育速率与温度的关系
Figure 1. Growth rate of L. invasa as affected by temperature
表 1 不同温度下桉树枝瘿姬小蜂各虫态的发育历期
Table 1. Durations of development stages of L. invasa under varied temperatures
虫态
Insect state发育历期 Developmental duration/d 19℃ 22℃ 25℃ 28℃ 31℃ 卵 Egg 10.33±1.56 aD 7.40±1.99 bD 6.33±1.73 cC 5.23±1.57 dC 4.33±1.24 eC 幼虫 Larva 66.13±4.58 aA 51.43±4.49 bA 43.27±4.65 cA 34.47±3.75 dA 28.17±3.21 eA 蛹 Pupa 19.63±2.41 aC 12.27±2.53 bC 9.37±1.77 cB 7.27±1.89 dB 5.40±1.30 eBC 产卵前期 Pre-oviposition 12.30±1.53 aD 5.13 ±2.34 bD 2.03 ±0.82 cD 1.05 ±0.52 dD 0.48 ±0.35 dD 成虫 Adult 23.97±3.23 aB 14.23±2.57 bB 10.33±1.63 cB 8.20±1.77 dB 6.27±1.57 eB 全世代 Generation 108.40±6.12 a 76.23±7.31 b 60.99±6.05 c 48.02±5.27 d 38.38±4.52 e 注：同行数据后不同小写字母表示同一阶段不同温度间差异显著，同列数据后不同大写字母表示同一温度不同世代间差异显著（SNK法, P＜0.05）；表中“成虫历期”指的是成虫寿命，世代历期=卵历期+幼虫历期+蛹历期+产卵前期，表2～4同。
Note: Different lowercase letters in the same row show significant difference, and different capital letters in the same column show significant difference（SNK, P＜0.05）；In the table, "adult duration" refers to adult life span；Generation duration = egg stage + larva stage + pupa stage + pre spawning stage. The same as Table 2–4.
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表 2 桉树枝瘿姬小蜂各虫态发育速率及成虫寿命与温度的关系模型
Table 2. Regression models on growth rate and adult lifespan of L. invasa as affected by temperature
虫态
Insect state图测法 Graph method 线性回归 Linear regression Logistic回归 Logistic regression 模型
Model决定系数（R2）
Determination coefficient模型
Model决定系数（R2）
Determination coefficient模型
Model决定系数（R2）
Determination coefficient卵 Egg v=0.011 5t−0.124 0.983 2 v=0.012 1t−0.128 0 0.488 9 v=0.366 4/（1+e3.554 7−0.138 4t） 0.487 7 幼虫 Larva v=0.001 7t−0.017 5 0.986 3 v=0.001 7t−0.018 5 0.833 6 v=0.154 8/（1+e3.834 6−0.149 5t） 0.843 8 蛹 Pupa v=0.010 8t−0.158 1 0.985 0 v=0.011 8t−0.176 7 0.682 5 v=0.461 2/（1+e4.659 8−0.189 6t） 0.688 7 产卵前期 Pre-oviposition v=0.273t-5.645 7 0.790 1 v=0.273t-5.645 7 0.349 4 v=0.898 6/（1+e2.709 5−0.109 3t） 0.497 3 成虫 Adult v=0.009 6t−0.141 4 0.994 7 v=0.010 4t−0.156 9 0.733 9 v=0.305 9/（1+e4.792 7−0.181 4t） 0.736 0 全世代 Generation v=0.001 4t−0.017 8 0.992 4 v=0.001 4t−0.017 8 0.898 1 v=0.684 8/（1+e3.145 1−0.131 9t） 0.901 1
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表 3 赣南桉树枝瘿姬小蜂各虫态的发育适温区范围
Table 3. Temperature ranges for optimal development at various stages of L. invasa in southern Jiangxi
虫态
Insect stateLogistic模型参数
Parameters of logistic model最适温Tmid
Optimum temperature/℃适温下限Tmin
Lower limit of temperature/℃适温上限Tmax
Upper limit of temperature/℃a b 卵 Egg 3.554 7 0.138 4 25.68 13.58 37.79 幼虫 Larva 3.834 6 0.149 5 25.65 13.85 37.45 蛹 Pupa 4.659 8 0.189 6 24.58 14.05 35.11 产卵前期 Pre-spawn 2.709 5 0.109 3 24.79 12.25 37.33 成虫 Adult 4.792 7 0.181 4 26.42 15.23 37.61 全世代 Generation 3.145 1 0.131 9 23.84 12.21 35.48
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表 4 桉树枝瘿姬小蜂发育历期预测值与观测值的相关性分析
Table 4. Predicted and observed durations of L. invasa development stages
虫态
Insect state观测值 Observed value/d 预测值 Predicted value/d 相关系数（r）
Correlation coefficientP值
P Value19℃ 22℃ 25℃ 28℃ 31℃ 19℃ 22℃ 25℃ 28℃ 31℃ 卵 Egg 10.33 7.40 6.33 5.23 4.13 10.58 7.75 6.12 5.05 4.30 0.995 8 0.000 3 幼虫 Larva 66.13 51.43 43.27 34.47 28.17 67.57 50.25 40.00 33.22 28.41 0.994 3 0.000 5 蛹 Pupa 19.63 12.27 9.37 7.27 5.40 21.23 12.58 8.94 6.93 5.66 0.997 4 0.000 2 全世代 Generation 108.40 76.25 60.99 48.02 38.38 113.64 76.92 58.14 46.73 39.06 0.997 5 0.000 2
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表 5 桉树枝瘿姬小蜂的LDT和SET （文献资料[10-13]及本试验）
Table 5. LDT and SET of L. invasa （data from literatures and this experiment）
地点
Position起点温度 LDT/℃ 有效积温 SET/（d·℃） 食料、温度
Food, Temperature卵
Egg幼虫
Larva蛹
Pupa成虫
Adult卵
Egg幼虫
Larva蛹
Pupa成虫
Adult福建福州
Fuzhou, Fujian 26.09°N8.65 6.57 13.94 14.83 101.80 807.16 105.43 114.53 巨桉8号, 21～36℃, 6个恒温[13]
E. grandis, 21～36℃, 6 constant temperatures江西赣州
Ganzhou, Jiangxi 25.85°N10.78 10.29 14.64 14.73 86.96 588.24 92.59 104.17 DH201-2, 19～31℃, 5个恒温(本研究结果)
DH201-2, 19～31℃, 5 constant temperatures [this experiment]广东广州
Guangzhou,Guangdong 23.08° N13.00 19.71 17.07 – 128.10 284.24 201.22 – 尾赤桉[12]
E. urophylla × E. camaldulensis广东广州
Guangzhou,Guangdong 23.08° N0.00 0.00 0.00 0.00 146.57 1228.01 161.67 205.17 DH201-2，室外，变温[11]
DH201-2, Outdoor, variable temperature广西南宁
Nanning, Guangxi 22.84°N10.25 10.79 749.43 53.42 20.5，27.5，28.6℃, 3个恒温[11]
20.5，27.5，28.6℃, 3 constant temperatures
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