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桃蛀果蛾外部形态和超微结构研究(鳞翅目:蛀果蛾科)

Morphology And Ultrastructure of The Peach Fruit Moth Carposina Sasakii Matsumura (Lepidoptera: Carposinidae)

【作者】 陈静

【导师】 花保祯;

【作者基本信息】 西北农林科技大学 , 农业昆虫与害虫防治, 2015, 博士

【摘要】 桃蛀果蛾Carposina sasakii Matsumura(鳞翅目Lepidoptera:蛀果蛾科Carposinidae)是东亚果园重要的蛀果害虫,以幼虫蛀食果实,寄主植物包括桃、杏、山楂、苹果、梨、枣、酸枣和石榴等10多种果树,常造成严重经济损失。目前对该虫的研究大多集中在生活史、种群动态和综合防治等方面,而对其形态学,特别是超微形态学鲜有涉及。本文借助光学显微镜、扫描电子显微镜和透射电子显微镜,研究了桃蛀果蛾成虫以及卵、幼虫、和蛹的形态及其相关感器,可望为该虫的某些行为提供形态学依据;并运用几何形态测量学分析不同寄主上桃蛀果蛾翅形的差异,为研究该虫的形态分化提供依据。主要研究结果如下:1.成虫形态:桃蛀果蛾成虫触角线状,复眼位于头部两侧。口器虹吸式,雌性下唇须明显长于雄性。前翅翅面近基部和中部有7–9簇黄褐色斜立鳞片丛,后翅灰白色。前足有净角器,中足和后足胫节有距。口器、触角和外生殖器上感器较为丰富。成虫口器主要由1个卷曲的喙管和1对发达的下唇须组成,其余部分均有不同程度退化。喙管由下颚的外颚叶演化而成。3节的下唇须表现出明显的性二型现象,雌性明显长于雄性。口器上有7种感器:刺形、鳞形、钟形、毛形、锥形、栓锥形和扁平感器。触角上共发现8种感器,分别为毛板形感器、栓锥感器、耳形感器、腔锥感器、刺形感器、锥形感器、鳞形感器和毛形感器。柄节和鞭节上分布有毛板形感器和鳞形感器,鞭节上除毛板形感器外其余感器均有分布。复眼由大量的小眼组成。每个小眼是由1个角膜、1个晶锥、9个视网膜细胞、1对初级色素细胞和6个次级色素细胞组成。8个视网膜细胞(R1-R8)的视小杆相互连接形成一个大的视杆,延伸至基膜,第9个视网膜细胞(R9)位于小眼基部。翅面鳞片按形状可分为4类,鳞片I扁平宽阔,端部不分叉,数量最多,主要分布在翅面最底层;鳞片II较狭长,端部亦不分叉,数量最少,主要分布在靠近翅的边缘区域;鳞片III扁平宽阔,覆盖在鳞片I上,少数成簇的斜立在翅面上,称为鳞片丛;鳞片IV最为狭长,其端部分4-5个叉,主要分布在翅的边缘。鳞片III和IV形状最为特殊,特别是部分鳞片III成簇斜立在翅面近基部和中部。雌性交配囊卵圆形,中央有1对“π”形交配囊片。前阴片基部有交配囊孔。肛突为一圆形瓣状结构,表面具有大量的刺形感器。雄蛾背兜较大,头巾状。抱器瓣内侧密被大量的毛,基内侧为粗长而弯的强齿突。阳茎针位于阳茎端膜末端,为大量的棒状结构。颚形突臂细长,上举。钩状突端部密生长刚毛。囊形突圆锥状。基于几何形态测量学不同寄主桃蛀果蛾翅形的分析:翅形普氏距离比较分析发现,东北梨上的桃蛀果蛾与其他寄主上的亲缘关系最远,这与典型变量(CVA)分析的结果一致。东北梨和陕西枣上的桃蛀果蛾翅的质心值较小,杏和苹果上的较大。2.卵壳超微形态:卵为椭圆形,顶端有4个卵孔。在卵孔周围有6–8瓣花饰小叶的莲花状花饰和2–3圈顶端分叉的“Y”形突起。卵壳侧面有规则排列的突起,突起交接处有1个凸出于卵表面的气孔。3.幼虫超微形态:桃蛀果蛾与山茱萸蛀果蛾幼虫触角和口器感器的类型和数量极为相似。二者触角第一节上均未发现任何感器;第二节具有3个锥形感器和2个刺形感器;第三节上有1个栓锥感器和3个锥形感器。口器上主要有6种感器,即刺形、锥形、栓锥形、指形、板形和内唇感器,区别在于山茱萸蛀果蛾比桃蛀果蛾少1对内唇感器。桃蛀果蛾老熟幼虫前胸上有10对刚毛,分别为前背毛、背毛、亚背毛、侧毛、亚腹毛,均为2对;中胸缺前背毛,侧毛3对,亚腹毛仅1对,其余与前胸一样。后胸和中胸的毛序极为相近。第1腹节背毛、亚背毛、亚腹毛均为2对,侧毛3对;第2腹节毛序与第1腹节毛序相近,只是亚腹毛有3对;第3–6腹节与第1、2腹节毛序相近,只是亚腹毛有4对;第7腹节背毛,亚背毛的分布均与前几腹节一致,侧毛2对,亚腹毛2对;第8腹节背毛、亚背毛均为2对,亚侧毛3对,1对亚腹毛;第9腹节毛序较为简单,背毛2对,亚背毛、侧毛和亚腹毛均为1对;臀板毛序最为简单,仅有2对背毛和2对亚背毛。桃蛀果蛾老熟幼虫趾钩为单序环状,3至6腹节上每个腹足趾钩为15–21个,臀足趾钩为9–13个。1龄幼虫腹足趾钩仅有4个,数量较少。4.蛹的形态:蛹为被蛹,长椭圆形,橙黄色,光滑。头部有红色的复眼、触角、上唇、上颚、喙管、下颚须及下唇须。前翅达到第4腹节腹面后缘。腹部10节。1–8腹节两侧各有1个气门。雌蛹生殖孔和产卵孔相连成一纵裂缝,位于第8和第9腹节腹面中央。雄蛹生殖孔位于第9腹节腹面中央。肛门位于第10腹节腹面。无臀棘。本研究对单食性的山茱萸蛀果蛾和多食性的桃蛀果蛾幼虫触角和口器的结构进行了对比分析,结果表明两种蛀果蛾幼虫触角和口器的感器与寄主范围之间并无严格的对应关系。通过对不同寄主上桃蛀果蛾翅形的量化分析,认为梨上的桃蛀果蛾与其它寄主上的桃蛀果蛾亲缘关系较远。

【Abstract】 The peach fruit moth, Carposina sasakii Matsumura(Lepidoptera: Carposinidae), is one of the serious pests of orchards in the eastern Asia, with the larvae boring in the fruits of various important fruit trees, frequently causing significant damage. Its host plants include several economically important fruits, such as Prunus persicae, P. armeniaca, Crataegus pinnatifida, Malus domestica, Pyrus spp., Zizyphus jujuba, Z. spinosus, and Punica granatum. Previous studies on C. sasakii mainly focused on its life cycle and integrated control, the morphology has not been well studied. In this thesis, the morphology of the adults, egg, larva and pupa in C. sasakii were investigated using light microscopy and scanning and transmission electron microscopy, in order to provide morphological basis for some behaviors. The wing shape of the C. sasakii on the different host plants was analysised using geometric morphometrics, to provide morphological basis for its morphological diferentitaion. The main results are as follows:1. Morphology of the adults: The antennae are filiform. The mouthparts are in the siphoning type, and the paired three-segmented labial palps of the female are longer than those of the male. There are 7–9 clusters of erect scales on the fore wing, obliquely. We found the epiphysis on the forelegs, medial spurs and apical spurs on the midlegs and hindlegs. Numerous sensilla are present on the mouthparts, antennae and genitalia.The mouthparts consist of a striking coilable proboscis formed from two galeae of the maxillae, a pair of developed labial palps, and other degenerate parts. The three-segmented labial palps display distinct sexual dimorphism between the sexes, much longer in the female than in the male. The mouthparts aslo bear seven types of sensilla, i.e. sensilla campaniformia, s. trichodea, s. styloconica, s. basiconica, s. squamiformia, s. chaetica and flattened sensilla.The surface of antennae has irregular reticular pattern, except the smooth base of the scape and pedicel. Eight types of sensilla are found on the antennae: hair plate sensilla, sensilla styloconica, s. auricillica, s. coeloconica, s. chaetica, s. basiconica, s. squamiformia and s. trichodea. The basal scape and the pedicel only possess hair plate sensilla and sensilla squamiformia. The terminal flagellum carries all sensilla types, except the hair plate sensilla.The compound eyes are composed of numerous ommatidia, each of which consists of a convex cornea, a crystalline cone, nine retinula cells, two primary pigment cells, and six secondary pigment cells. The rhabdom is formed by the rhabdomeres from eight retinula cells and runs to the basal lamina. The 9th retinula cell only lies on the basal lamina.Four types of scales are found on the wing. Type I is flatted and broad, forms forked branch. The scales are abundant and widespread on the bottom of the wing. Type II is slender than type I, does not form forked branch. The scales are the fewest and are distributed near the wing margin. Type III are also flatted and broad, covered on scales I. A few of type III are clustered and oblique on the wing, and referred to as “scale tufts”. Type IV is the longest and forked, and distributed on the wing margin. A few of type III are cluster and oblique at the base and middle in the wing.In the female, a pair of π-shaped signa is found on the middle of the oval corpus bursae. The ostium bursa is found on the venter of lamella antevaginalis. The papillae anales are oval and valvular structures, and have many sensilla chaetica on their surface.In the male, the tegumen is very large and turban-shaped. The interior valvae have many setae, and have strong and long tooth. Cornuti are found on the distal end of vesica. The gnathos arms are long and upward. The top of uncus have many long setae. Saccus is conical.Geometric morphometric analysis of wing shape on different host plants: In comparison with other host plants, the procustues distance of C. sasakii is the farthest on Pyrus spp. The result is in accordance with that of canonical variance analysis. On the comparision trials of the centroid sizes on wing shape, C. sasakii are smaller on Pyrus spp. and Z. jujuba, and are larger on P. armeniaca and M. domestica.2. Ultrastucture of the eggshell: The eggs of C. sasakii are oval, with the micropyles located at the top of the egg. The micropyles are usually four in number. Lotus-like decoration with 6–8 disc floriation lobules and 2–3 circles of Y-shaped processes are around the micropyles. The granular processes and spiracles are located on the lateral side of eggs.3. Ultrastucture of the larva: The types and numbers of sensilla on the larval antennae and mouthparts of C. sasakii and C. coreana are very similar. The first segment of the antenna is devoid of sensilla. The second segment possesses two sensilla chaetica and three sensilla basiconica. The terminal segment carries distally one sensillum styloconicum and three sensilla basiconica. Six types of sensilla are found on the mouthparts: sensilla chaetica, s. styloconica, s. digitiformia, s. basiconica, s. placodea and epipharyngeal sensilla. C. sasakii has three pairs of epipharyngeal sensilla, and C. coreana has two pairs of epipharyngeal sensilla.There are ten pairs of setae on the prothorax of mature larvae: prothoracic setae, dorsal setae, lateral setae, subdorsal setae, and subventral setae. There are no prothoracic setae, and three pairs of lateral setae and one pair of subventral setae on the mesothorax. The remaining setae on the mesothorax are similar to those of prothorax. The setae on the metathorax are similar to those on the mesothorax. There are two pairs of dorsal setae, subdorsal setae and subventral setae respectively, and three pairs of lateral setae on the first abdominal segment(A1). The setae on A2 are similar to the setae on A1, but three pairs of subventral setae. The setae on A3 to A6 are similar to the setae on A1 and A2, but four pairs of subventral setae. There are two pairs of dorsal setae, subdorsal setae, lateral setae and subventral setae on A7, respectively. Two pairs of dorsal setae, subdorsal setae, three pairs of lateral setae and one pair of subventral setae are found on A8. Two pairs of dorsal setae, one pair of subdural setae, lateral setae and subventral setae are found on A9. Only two pairs of dorsal setae and subdorsal setae are on the anal plate.The crochets of the mature larvae are in a uniordinal circle. There are 15–21 crochets on each abdominal proleg from A3 to A6, and 9–13 crochets on A11. The crochets of the first instar are only 4 in number.4. Morphology of the pupa: The pupa of C. sasakii is obtect, glabrous and orange yellow. Its head includes the red compound eyes, antennae, labrum, mandibles, maxillary palps, proboscis, labial palps. The wing-pad of fore wing reaches the 4th abdominal segment. The pupal abdomen consists of ten segments, the last three of which are always fused together. There is a spiracle on both sides of A1 to A8. In male pupae the genital groove occurs midventrally on A9, and in female genital grooves are usually found midventrally on A8 and A9 and confluent longitudinal crack. The anus is situated on A10. No cremaster was found.Through morphological comparisons of sensilla on the larval antennae and mouthparts of C. sasakii and C. coreana, we concluded that there is no strict correspondence between larval sensilla and host range in Carposinidae. The wing shape analysis of C. sasakii on the different host plants indicates that the genetic divergence of C. sasakii is the greatest on Pyrus spp. compared with other host races.

【关键词】 桃蛀果蛾山茱萸蛀果蛾幼虫成虫口器触角复眼鳞片外生殖器
【Key words】 Carposina sasakiiCarposina coreanaegglarvapupaadultmouthpartsantennacompound eyescalegenitalia
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