椰子织蛾幼虫龄数及取食量的雌雄差异
2017-09-16林玉英金启安温海波唐雅文彭正强
林玉英,金 涛,金启安,温海波,唐雅文,彭正强
(中国热带农业科学院环境与植物保护研究所,农业部热带农林有害生物入侵监测与控制重点实验室,海南儋州 571737)
椰子织蛾幼虫龄数及取食量的雌雄差异
林玉英,金 涛,金启安,温海波,唐雅文,彭正强*
(中国热带农业科学院环境与植物保护研究所,农业部热带农林有害生物入侵监测与控制重点实验室,海南儋州 571737)
为了明确椰子织蛾幼虫的龄数、取食量及龄期。在室温25℃±3℃,寄主食料椰子叶饲养条件下,测量了雌、雄幼虫头壳宽、取食量并记录了各龄幼虫的发育历期。结果表明,椰子织蛾雌性幼虫有9-10个龄数,雄性幼虫有8-10个龄数。1-10龄雌虫的头壳宽约为0.2300,0.3250,0.4300,0.5267,0.7700,0.9633,1.3775,1.5850,1.8200,2.1929 mm。1-10龄雄虫的头壳宽约为0.2233,0.3214,0.4125,0.5300,0.6529,0.8675,1.1267,1.3375,1.4950,1.8925 mm。9-10龄雌虫的头壳宽显著大于雄虫的头壳宽。头壳宽与龄数具有很强的相关性。椰子织蛾幼虫的取食量随龄数的增大而增加。椰子织蛾1代雌性幼虫平均取食椰子叶的面积(3607.23±146.83 mm2)显著高于1代雄性幼虫平均取食椰子叶的面积(1991.25±143.92 mm2)。1-5龄幼虫的取食量最小,小于50 mm2。8-10龄为暴食期。1-10龄幼虫的发育历期分别为4.55±0.16,5.69±0.24,5.73±0.37,5.22±0.15,5.11±0.46,4.61±0.46,5.12±0.68,6.00±0.43,6.86±0.40和8.75±1.55 d。对于雌、雄成虫个体差异较大的昆虫,对其幼虫头壳宽值和取食量的测定应雌、雄幼虫分别测定。椰子织蛾的防治适期应在未造成严重为害的1-5龄幼虫高蜂期进行。以幼虫头壳宽为主,同时结合取食面积判定幼虫所处龄数,可为准确掌握防治时期提供科学依据。
椰子织蛾;性别;龄数;取食量;龄期
椰子织蛾OpisinaarenosellaWalker是棕榈科植物上的重要害虫,2014年被国家林业局列入外来入侵有害生物名单。寄主植物包括椰子Cocosnucifera、中东海枣Phoenixsylvestris、大王棕Roystonearegia(HBK.)O.F. Cook、槟榔Arecacatechu、蒲葵Livistonachinensis(Jacq.)R. Br、华盛顿棕Washingtoniarobusta、散尾葵ChrysalidocarpuslutescensH. Wendl.、桄榔Arengapinnata(Wurmb.)Merr.、圆叶轴榈LicualagrandisH.Wendl.、红脉榈Latanialontaroides、斐济榈PritchardiaPacifica、香蕉MusananaLour.、甘蔗Saccharumofficinarum等(吕宝乾等,2013)。该虫已在印度、泰国、缅甸、印度尼西亚等东南亚国家和地区发生,并对当地棕榈科植物造成重大经济损失(Jayaratnam,1941;吕宝乾等,2013)。椰子是椰子织蛾最喜食的寄主,危害严重时可将超过90%的椰子叶片吃光,是印度和斯里兰卡椰子树上的毁灭性害虫(Nirula,1956;阎伟等,2013)。该虫的危害致使椰子落果率增大,植株发育缓慢,严重时致使椰子产量降低50%以上(Remachandran,1979)。幼虫在叶背面形成不规则蛀道,蛀道内粪便与其吐的丝交织,幼虫隐藏于蛀道内取食叶肉,严重时叶肉被吃光,叶片卷折、干枯。在海南,椰子织蛾完成1个世代需2-2.5个月,1年发生5代,世代重叠严重(李洪等,2015)。野外终年可见椰子织蛾各种虫态,幼虫正常取食,卵、蛹正常发育,成虫均可产卵,无越冬越夏现象。目前,海南已有9个市县发现椰子织蛾危害,并已在广东、福建和广西省发现该虫危害。
昆虫学工作者通常测定幼虫头壳宽度,以头宽作为划分幼虫龄数的标准(束春娥和曹赤阳,1989;祁诚进,1991)。吕宝乾等(2013)报道了椰子织蛾幼虫有5-8个龄数;Perera等(1988)报道了椰子织蛾幼虫通常有5个龄数;Babu 和Prabhu (1989)报道了椰子织蛾在实验室条件下分为8龄。由于椰子织蛾幼虫龄数报道的不一致,为准确掌握和预测该虫的发生趋势,本试验测定椰子织蛾各龄幼虫的头壳宽度和取食量,并记录其发育历期。在利用天敌昆虫开展生物防治的过程中,需要根据各龄幼虫的发生情况来释放天敌。同时从被取食叶面积的大小可以推测幼虫龄数,取食量的测定可为确定防治适期提供依据。幼虫龄数、龄期和取食量的研究有助于生命表及预测预报和防治的研究(刘秀琼和曾仁光,1981)。
1 材料与方法
1.1 供试虫源
椰子织蛾采自海南省儋州市,采集野外被椰子织蛾为害的椰子叶,置于室内饲养,待成虫羽化后,收集于产卵笼中,提供棉花团和新鲜的椰子老叶供其产卵,并以10%蜂蜜水供其补充营养,次日将带卵的棉花团和椰子叶取出,待卵孵化后,取12 h内的初孵幼虫供试。饲养环境条件:室内温度25℃±3℃。
1.2 方法
1.2.1各龄幼虫头壳宽和取食量的测量及发育历期的记录
将椰子老叶剪成6 cm长置于直径3 cm,高8 cm 玻璃管中,将初孵幼虫单头饲养于玻璃管中并编号,用棉花团塞住管口,3天更换1次椰子叶,每天定时观察幼虫是否蜕皮并记录蜕皮时间,饲养至成虫阶段,记录羽化成虫的性别。将1-3龄幼虫取食的叶片部位置于Carl zeiss SteREO Discovery型体视显微镜下,调至图像清晰后,使用Axio Vision软件拍照后用Outline工具进行测量,4-10龄幼虫取食的叶面积利用透明坐标纸估算。用显微镜测微尺测量幼虫所蜕下来头壳的宽度(陈湖,1990)。参试幼虫50头。
1.2.2数据分析
各龄幼虫指标之间的差异统计采用SPSS软件的单因素方差分析进行Duncan多重比较,雌雄之间的指标差异采用t检验法。对头壳宽与龄数间的关系利用SPSS软件和Excel软件进行多项式回归分析。总取食面积为每头幼虫各龄期取食量之和。
2 结果与分析
2.1 椰子织蛾幼虫龄数和头壳宽
用椰子叶饲养的椰子织蛾雌性幼虫有9-10个龄数,雄性幼虫有8-10个龄数。头宽随龄数的增加而增大(表1)。雌虫头壳宽为0.2300-2.1929 mm,雄虫头壳宽为0.2233-1.8925 mm。1-10龄雌虫的头壳宽为0.2300,0.3250,0.4300,0.5267,0.7700,0.9633,1.3775,1.5850,1.8200,2.1929 mm。1-10龄雄虫的头壳宽为0.2233,0.3214,0.4125,0.5300,0.6529,0.8675,1.1267,1.3375,1.4950,1.8925 mm。1、2、3龄雌虫头壳宽差异不显著(P>0.05),4-10龄雌虫头壳宽差异显著(P<0.05)。1-5龄雄虫前后两龄头壳宽差异不显著(P>0.05);8龄和9龄雄虫头壳差异不显著(P>0.05);6龄、7龄、10龄雄虫与其他龄数雄虫头壳宽差异显著(P<0.05)。1-10龄雌性、雄性幼虫的头壳宽值没有相互重叠。9-10龄雌虫的头壳宽显著大于雄虫的头壳宽(P<0.05)。前后两龄头壳宽均值比在1.12-1.46,基本符合戴氏定律(Dyar’s rule)。
表1 椰子织蛾各龄幼虫的头壳宽
注:括号外面的字母是不同龄数雌虫、雄虫头壳宽值之间邓肯氏新复极差多重比较的结果,括号里面的字母是同一龄数雌虫、雄虫头壳宽值之间多重比较(t检验)的结果,相同字母表示差异不显著(P>0.05)。Note: Same letters outside of the brackets represent no significant difference among head capsule widths of different instars of female or male larvae (P>0.05; DMRT). Same letters inside brackets show no significant difference between head capsule widths of female and male larvae of the same instar (P>0.05; t-test).
椰子织蛾幼虫的头宽随幼虫龄数的增加呈上升趋势(图1)。经拟合分析,头壳宽与龄数具有很强的相关性;雌性幼虫头壳宽与龄数间的关系符合多项式方程:y=0.015x2+0.048x+0.147,(R2=0.994,df=2,7,F=595.987,P<0.0001);雄性幼虫头壳宽与龄数间的关系符合多项式方程:y=0.013x2+0.034x+0.184,(R2=0.995,df=2,7,F=828.371,P<0.0001)。5-10龄雌虫的头壳宽大于雄虫的头壳宽。
2.2 椰子织蛾雌、雄幼虫的取食量和为害状
椰子织蛾雌、雄幼虫的取食量随着龄数的增大而增加(表2)。雌性10龄幼虫的取食量(1800.75±82.75 mm2)和9龄的(1810.7±58.68 mm2)差异不显著(P>0.05);10龄和9龄的取食量显著大于8龄的(889.41±80.37 mm2)(P<0.05);8龄的取食量显著大于7龄的(210.00±16.78 mm2)和6龄的(114.70±6.39 mm2),并显著大于1-5龄的(0.36-43.40 mm2)(P<0.05);整个幼虫期每头雌性幼虫可累计取食椰子叶片面积为3607.23±146.83 mm2。雄性10龄幼虫的取食量(1351.33±200.60 mm2)显著大于9龄的(787.83±176.93 mm2)和8龄的(665.05±103.08 mm2)(P<0.05);9龄的和8龄的取食量显著大于7龄的(258.70±34.23 mm2)和6龄的(104.80±15.87 mm2),并显著大于1-5龄的(0.56-38.85 mm2)(P<0.05)。1龄雌性幼虫的取食量显著小于1龄雄性幼虫的取食量(P<0.05);9龄雌性幼虫的取食量显著大于9龄雄性幼虫的取食量(P<0.05);2-8龄和10龄的雌、雄幼虫取食量差异不显著(P>0.05)。整个幼虫期每头雄性幼虫可累计取食椰子叶片面积为1991.25±143.92 mm2,显著小于每头雌性幼虫整个幼虫期取食的叶片面积(3607.23±146.83 mm2)(P<0.05)。椰子织蛾1-5龄幼虫的取食量最小,小于50 mm2。椰子织蛾1-2龄幼虫未啃食完取食处的叶肉,沿叶脉取食,留下叶脉。3-10龄幼虫啃食完取食处的椰子叶肉,留下下表皮,取食斑随龄数的增长而增大。8-10龄为暴食期。椰子织蛾为害严重时将叶肉吃光,使叶片形成干枯状。
图1 椰子织蛾雌、雄幼虫头壳宽与龄数模拟模型Fig.1 Model for head capsule widths of female and male Opisina arenosella larvae with different instars
表2 椰子织蛾雌、雄幼虫取食量
注:括号外面的字母是不同龄数雌虫、雄虫取食量之间邓肯氏新复极差多重比较的结果,括号里面的字母是同一龄数雌虫、雄虫取食量之间多重比较(t检验)的结果,相同字母表示差异不显著(P>0.05)。Note: Same letters outside of the brackets represent no significant difference among feeding amounts of different instars of female or male larvae (P>0.05; DMRT). Same letters inside brackets show no significant difference between feeding amounts of female and male larvae of the same instar (P>0.05; t-test).
2.3 椰子织蛾幼虫发育历期
椰子织蛾幼虫发育历期最短为39 d,最长为60 d,平均需要48.85±7.06 d(表3)。各龄幼虫的发育历期最短为4 d,最长为13 d。1-10龄幼虫的发育历期分别为4.55±0.16,5.69±0.24,5.73±0.37,5.22±0.15,5.11±0.46,4.61±0.46,5.12±0.68,6.00±0.43,6.86±0.40和8.75±1.55 d。1龄幼虫的龄期最短,10龄幼虫的龄期最长。1-8龄幼虫的龄期差异不显著(P>0.05),并显著短于10龄幼虫的龄期(P<0.05);1龄,5龄和6龄幼虫的龄期显著短于9龄和10龄幼虫的龄期(P<0.05);10龄幼虫的龄期显著长于1-9龄幼虫的龄期(P<0.05)。
表3 椰子织蛾幼虫各龄发育历期
注:字母是不同龄数椰子织蛾幼虫龄期之间邓肯氏新复极差多重比较的结果,相同字母表示差异不显著(P>0.05)。Note: Same letters represent no significant difference among durations of different instars ofO.arenosellalarvae (P>0.05; DMRT).
3 结论与讨论
幼虫龄数划分是生物学研究的基础之一。椰子织蛾雌性幼虫有9-10个龄数,雄性幼虫有8-10个龄数。可能是由于获得有效营养不均、温度、自然光照周期、叶片剪下数天后营养成份变化,或者由于实验室的饲养条件不完全适于昆虫的生长发育,幼虫生长受到干扰,从而造成椰子织蛾幼虫龄数不一致。温度能够影响幼虫的取食行为从而影响幼虫的龄数(Delbacetal., 2010)。Gaines和Campbell(1935)发现美洲棉铃虫HeliothiszeaBoddie幼虫的龄数受食料的影响。Peterson和Haessler(1928)发现梨小食心虫GrapholithamolestaBusck幼虫的龄数受食料和温度的影响。根据王凤等(2009)报道可能是由于室内饲养环境造成幼虫获得有效营养不均,红棕象甲Rhynchophorusferrugineus幼虫有7-9个龄数。室内饲养导致了额外的幼虫龄数(Pereraetal., 1988)。本试验主要在2-3月份进行,自然界温度较低、光照时间较短可能是导致用椰子叶饲养的椰子织蛾雌性幼虫有9-10个龄数,雄性幼虫有8-10个龄数,多于前人报道的5-8个龄数(吕宝乾等,2013)的原因,本推测与Argyro(1996)报道的在短光照条件(10 L ∶14 D)下西非蛀茎夜蛾Sesamianonagrioodes有额外的龄数相一致。在本研究中1、2、3龄椰子织蛾雌虫头壳宽差异不显著;1-5龄雄虫前后两龄头壳宽差异不显著;存在不同龄数的幼虫头宽值差异不显著的现象。椰子织蛾幼虫1-3龄头壳增长较慢,约为0.1 mm/龄,还存在少数幼虫在蜕皮前后头宽值变化不明显的现象,如有1头雄性幼虫5龄期间头壳只增长了约0.01 mm,1-9龄头壳由0.22 mm增长到1.29 mm,到末龄(10龄)快速增长至1.92 mm。椰子织蛾前后两龄幼虫头壳宽测量均值比在1.12-1.46,这一结果基本符合戴氏定律(Dyar’s rule)——昆虫前后2龄头壳宽比值在1.2-1.4(彩万志和庞雄飞,2001)。椰子织蛾雌成虫一般明显大于雄成虫,5-10龄雌虫的头壳宽大于雄虫的头壳宽,因此对于雌、雄成虫个体差异较大的昆虫,对其幼虫头壳宽值的测定应雌、雄幼虫分别测定。椰子织蛾各龄幼虫的头壳宽度值不重叠,且与龄数具有很强的相关性,头壳宽值变异较小,可作为识别幼虫龄数的主要特征。马素芳(1964)报道用头的宽度辨别不同龄幼虫是比较可靠的。椰子织蛾幼虫取食量随着龄数的增大而增加,应用硬化的头壳作为区分椰子织蛾幼虫龄数的主要依据,同时结合取食斑的面积可以快速准确鉴定幼虫龄数,确定田间种群发育所处的虫龄阶段,为该虫的预测预报以及确定有效防治时期提供科学依据(陈永年和潘桐,1988;钟义海等,2005;王春蕾等,2007)。
一般1龄幼虫很快,其龄期与其他龄数相比相应较短;而末龄幼虫则需要更长时间完成发育(王小艺等,2012;王雪龙,2015),进入预蛹和蛹期前经历暴食到逐渐停止取食,排空体腔这一过程。本研究中椰子织蛾1龄幼虫的龄期最短,10龄幼虫的龄期最长,10龄幼虫的龄期显著长于1-9龄幼虫的龄期,进入预蛹前的8-10龄为暴食期,说明该虫也符合这一规律。
椰子织蛾雌性幼虫的取食量与雄性幼虫的取食量存在差异,椰子织蛾1代雌性幼虫平均取食椰子叶的面积(3607.23±146.83 mm2)显著高于1代雄性幼虫平均取食椰子叶的面积(1991.25±143.92 mm2),这是椰子织蛾雌成虫一般显著大于雄成虫的主要原因,因此对于雌、雄成虫个体差异较大,对其幼虫取食量的测定应雌、雄幼虫分别测定。有文献报道三星黄萤叶甲Parideaangulicollis雌虫成虫取食量显著大于雄成虫的取食量(宋佃远,2007)。茄二十八星瓢虫Henosepilachnavigintioctopuctata雌成虫取食量显著大于雄成虫的取食量(杨晨亮等,2011)。
8-10龄椰子织蛾幼虫进入暴食期,短时间内取食大面积叶片,使椰子叶大面积干枯,此时期危害最严重;椰子织蛾的防治适期应定在未造成严重为害的1-5龄幼虫高蜂期进行,即田间被害寄主叶片被幼虫取食面积为50 mm2以下时施药,可有效地控制其为害。取食量的测定为制定椰子织蛾防治指标提供了依据。
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DistinctionininstarsandfeedingamountsbetweenOpisinaarenosellaWalkerfemaleandmalelarvae
LIN Yu-Ying, JIN Tao, JIN Qi-An, WEN Hai-Bo, TANG Ya-Wen, PENG Zheng-Qiang*
(Institute of Plant and Environment Protection, Academy of Tropical Agriculture Sciences of China, Ministry of Agriculture Key Laboratory for Monitoring and Control of Tropical Agricultural and Forest Invasive Alien Pests, Danzhou 571737, Hainan Province, China)
In order to define the instar numbers, feeding amount and duration of the instar ofOpisinaarenosellaWalker larva, the head capsule widths and feeding amounts of different instars ofO.arenosellafemale and male larvea fed on coconut leaves were tested, and durations of the instar were recorded under 25℃±3℃ in the laboratory. The results showed that female larvae had 9-10 instars and male larvae had 8-10 instars. The head capsule widths of 1st-10thinstar female larvae were about 0.2300, 0.3250, 0.4300, 0.5267, 0.7700, 0.9633, 1.3775, 1.5850, 1.8200, 2.1929 mm, respectively, and the head capsule widths of 1st-10thinstar male larvae were about 0.2233, 0.3214, 0.4125, 0.5300, 0.6529, 0.8675, 1.1267, 1.3375, 1.4950, 1.8925 mm, respectively. The head capsule widths of 9th-10thinstar female larvae were significantly wider than those of 9th-10thinstar male larvae. The head capsule widths were significantly correlated with the instar. Larval feeding amounts increased with an increase in the instar. The feeding amount ofO.arenosellafemale during larval stage (3607.23±146.83 mm2)was significantly more than that of male (1991.25±143.92 mm2). The feeding amounts of 1st-, 2nd-, 3rd-, 4th-, and 5th-instar larvae were the lowest (<50 mm2). The greedily eating period of the larva were 8th-10thinstars. The average duration of 1st-10thinstars were 4.55±0.16, 5.69±0.24, 5.73±0.37, 5.22±0.15, 5.11±0.46, 4.61±0.46, 5.12±0.68, 6.00±0.43, 6.86±0.40 and 8.75±1.55 d, respectively. If the individual differences for female and male insects are big, the head capsule widths and feeding amounts of female and male larvae should be tested separately. The optimum control stage ofO.arenosellalarvae is the period when larvae are mainly in 1st-5thinstars. Discrimination of the instar mainly according to the head capsule widths, and meanwhile, feeding amounts may provide scientific bases for accurate grasping preventing and controlling period.
OpisinaarenosellaWalker; sex; instars; feeding amount; instar duration
中央级公益性科研院所基本科研业务费 (2014hzs1J001);海南省自然科学基金 (20153067);国家支撑项目 (2015BAD08B03) ;公益性行业( 农业) 科研专项( 201403075);国家重点研发计划(2016YFC1201200)
林玉英,女,1984年生,福建莆田人,硕士,研究方向为害虫综合治理,E-mail:linyuying2008@163.com
*通讯作者Author for correspondce, E-mail:lypzhq@163.com
Received: 2016-09-01; 接受日期Accepted: 2017-02-17
Q965;S433
:A
1674-0858(2017)04-0912-07
林玉英,金涛,金启安,等.椰子织蛾幼虫龄数及取食量的雌雄差异[J].环境昆虫学报,2017,39(4):912-918.