若干种天牛幼虫形态结构与生活习性的适应关系
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摘要
天牛是重要的林木蛀干害虫,以幼虫钻蛀为害植物的组织。国内外众多学者已对天牛形态、生活史、生活习性、空间分布型和消化酶等方面做了大量研究,但鲜有人探讨天牛幼虫的形态与其危害习性的关系。本研究侧重天牛幼虫的形态特征及与幼虫取食习性(包括主要取食部位、寄主树种类群、是否排粪等)的相互关系,重要结果如下：
l、天牛幼虫外部形态特征与取食习性的关系
通过比较天牛3亚科15种天牛幼虫上颚形态,探讨其中9种老熟幼虫上颚发达程度与取食树种材质、部位的关系；比较16种天牛幼虫行动器官特征及与其排粪习性的关系,分析9种老熟幼虫气门与其排粪习性的关系。
基本明确天牛亚科7种幼虫上颚均为圆凿型,沟胫天牛7种幼虫和幽天牛亚科的1种幼虫为楔型。从上颚端部的形态看,圆凿型上颚,端部背面和内面具数条横沟；楔型上颚,端部背面具数条横沟,内面常具隆脊。9种老熟幼虫中,楔型上颚的天牛,外露上颚占头部面积比例越大,上颚越发达；上颚为圆凿型的天牛,外露上颚占头部面积比例越小,其上颚越发达,取食能力越强。主要取食阔叶树的幼虫上颚发达程度强于主要取食针叶树的幼虫上颚,主要在木质部中危害的幼虫上颚发达程度强于主要在韧皮部中危害的幼虫上颚。
观察的16利一天牛幼虫前胸背板翼瓣发达,具明显侧沟和上侧片,背中线前半明显。沟胫天牛亚科多具后背板褶,松幽天牛亚科和天牛亚科多不具后背板褶。及时将粪便排出的幼虫前胸背板后半部多具“凸”字纹、粗颗粒或刻点,而未及时排出粪便的种类则多具细皱纹,微粒或小圆点。及时排粪类天牛幼虫的步泡突较为发达,表现为多具瘤突,且瘤突粗大,纵侧沟明显；不及时排粪类天牛幼虫运动能力相对较弱,表现为多无痛突,具细皱纹,侧纵沟浅。
天牛幼虫9对气门均呈长椭圆形或卵形。其中,中胸气门面积最大,其次是腹部末端气门面积,均显著大于其它7对气门面积比例值。排粪类天牛各体节气门面积占侧面积比例值显著大于不排粪类幼虫。
2、天牛内部形态特征与取食习性的关系
通过比较分析9种老熟天牛幼虫消化道特征与其取食树种的关系,以及呼吸系统特点与其排粪习性的关系；测量4种老熟幼虫呼吸速率与其排粪特征的关系。
明确天牛幼虫消化道结构简单,消化道长度显著长于体长。其中,中肠最长,占整个消化道长度的0.63-0.73；后肠次之,占0.22-0.35；前肠最短,不足消化道长度的0.06。危害针叶树的幼虫前肠占消化道总长比例均低于危害阔叶树的幼虫前肠比例。
9种天牛老熟幼虫呼吸系统结构特点为,体内有两条主气管,分布于两侧。气管的前端连接前胸,后端延伸至腹部最末端。气管与两侧气门相连但后胸气门不发达,仅具痕迹。气管与气门相连之处分出的二级、三级气管及微气管,与复眼、触角、消化道等器官相连。排粪类天牛幼虫一级气管直径大于不排粪幼虫直径。排粪类天牛幼虫二级气管数量较多,且直径大于不排粪类幼虫。
4种老熟幼虫,排粪类的桑天牛Apriona germari和松墨天牛Monochamus alternatus的幼虫代谢速率显著大于不排粪类的双条杉天牛Semanotus bifasciatus和红缘天牛Asias halodendri幼虫。上述4种天牛幼虫的呼吸墒值差异不大,但排粪幼虫呼吸墒值还是大于不排粪类幼虫。
总之,上述结果表明,天牛幼虫的形态结构与其取食习性存在明显的适应关系。
Cerambycinae which are important forest boring beetles, caused enormous losses on China's forestry by damaging plant tissues. A lot of scholars from home and abroad have done much work in Cerambycinae's characteristics, life history, life habits, spatial distribution pattern and digestive enzymes, but few people investigated the adaptability of morphological structures and harming habits of cerambycid larvae. The study observed the cerambycid larvae's morphological structures and the relationship with feeding habits, which included the main feeding site, the material of host plants and timely discharged faeces and sawdusts or not. The main results are as follow:
1、The relationship of external morphological structures and feeding habits:observed the mandible of15cerambycid species belong to3subfamilies of Cerambycidae and investigated the relationship of9mature larvae's developed degree of mandible and the material of host plants or the main feeding site; compared the relationship of the characteristics of the moving organs of16cerambycid larvae and discharging habits, and analyzed the relationship of9mature larvae's spiracles and discharging habits.
The results showed that the7larvae of subfamily Cerambyciae had gouge-shaped mandible, and the7larvae of subfamily Lamiinae and the1larvae of subfamily Aseminae had wedge-shaped mandible. From the morphology of the mandible's top, the back and inner surface of gouge-shaped mandible had several transverse furrows; the back of wedge-shaped mandible had several transverse furrows and its'inner surface had strong oncus. In the9species of mature larvae, For the larvae with gouge-shaped mandible, the smaller of the area ratio of its exerted-visible mandible to head, the stronger its mandible is. For the larvae with wedge-shaped mandible, the larger of the area ratio of its exerted-visible mandible to head, the stronger its mandible is. The mandible of larvae feeding broadleaf were stronger than the mandible of larvae feeding conifer, and the mandible of larvae mainly harming xylem were stronger than the mandible of larvae mainly harming phloem.
Observation of16species of larvae, the alar lobe of pronotum was strong, and had obvious lateral furrow, epipleurun and the first half of median line. Most of the larvae of subfamily Lamiinae had postnotum fold, the larvae of subfamily of Cerambyciae and Aseminae did not. The latter half of pronotum in the larvae which timely discharged faeces and sawdusts had'convex' pattern and tough particles or punctum, and which didn't timely discharge faeces and sawdusts had fine wrinkles and particles or small round spot. The ambulatory ampullae in the larvae which timely discharged faeces and sawdusts were more developed, with tough and big verrucae. The sports ability in the larvae which did not timely discharged faeces and sawdusts were worse, with fine wrinkle and small verrucae.
All the spiracles of cerambycid larvae were long oval or oval. Among them, the relative area of mesostigma was the largest, followed by the area of abdominal end spiracles, were significantly greater than the other7on the spiracles area ratio.the relative area of spiracles in the larvae which timely discharged faeces and sawdusts were larger than the larvae which did not.
2、The relationship of the inside morphological structures and feeding habits:observed the characteristic of alimentary canal of9mature cerambycid larvae and the relationship of the material of host plants, investigated the characteristic of the respiratory systems of9mature cerambycid larvae and the relationship of discharging habits, measured the respiration rate of4mature cerambycid larvae and the relationship of discharging faeces and sawdusts.
The structure of larvae's alimentary canal was simple, and its'length was obviously longer than the body. Among them, the midgut was the longest, accouting for the alimentary canal length0.63-0.73; hindgut followed by accounting for0.22-0.35; foregut was the shortest, less than0.06of the whole length. For the larvae feeding conifer, its'ratio of foregut to alimentary canal was smaller than the larvae feeding broadleaf.
The characteristic of the respiratory systems of9mature larvae; the body had two main trachea, distributed in both sides. The front of trachea connected to the prothorax, and the back-end extended to the end of the abdominal. The trachea connected spiracles on both sides, but metastigma was not developed, with only trace. The connection point of trachea and spiracles separated the secondary trachea, thirdary trachea and tracheole, and connected to compound eyes, antenna, alimentary canal and other organs. The primary trachea's diameter of larvae timely discharged faeces and sawdusts was larger than the larvae did not. The larvae timely discharged faeces and sawdusts had a larger number of secondary trachea, and its'diameter were larger than the larvae did not timely discharged.
In the4mature larvae, the larvae which timely discharged faeces and sawdusts-the metabolic rate of Apriona germari and Monochamus alternates were larger than the larvae did not timely discharged-Semanotus bifasciatus and Asias halodendri. The respiratory entropy of4species of cerambycid larvae were not significantly different, but respiratory entropy of larvae timely discharged was larger than the larvae did not timely discharged.
In all, the results above showed that the cerambycid larvae's morphological structures and feeding habits obviously had adaptive relationship.
l、天牛幼虫外部形态特征与取食习性的关系
通过比较天牛3亚科15种天牛幼虫上颚形态,探讨其中9种老熟幼虫上颚发达程度与取食树种材质、部位的关系；比较16种天牛幼虫行动器官特征及与其排粪习性的关系,分析9种老熟幼虫气门与其排粪习性的关系。
基本明确天牛亚科7种幼虫上颚均为圆凿型,沟胫天牛7种幼虫和幽天牛亚科的1种幼虫为楔型。从上颚端部的形态看,圆凿型上颚,端部背面和内面具数条横沟；楔型上颚,端部背面具数条横沟,内面常具隆脊。9种老熟幼虫中,楔型上颚的天牛,外露上颚占头部面积比例越大,上颚越发达；上颚为圆凿型的天牛,外露上颚占头部面积比例越小,其上颚越发达,取食能力越强。主要取食阔叶树的幼虫上颚发达程度强于主要取食针叶树的幼虫上颚,主要在木质部中危害的幼虫上颚发达程度强于主要在韧皮部中危害的幼虫上颚。
观察的16利一天牛幼虫前胸背板翼瓣发达,具明显侧沟和上侧片,背中线前半明显。沟胫天牛亚科多具后背板褶,松幽天牛亚科和天牛亚科多不具后背板褶。及时将粪便排出的幼虫前胸背板后半部多具“凸”字纹、粗颗粒或刻点,而未及时排出粪便的种类则多具细皱纹,微粒或小圆点。及时排粪类天牛幼虫的步泡突较为发达,表现为多具瘤突,且瘤突粗大,纵侧沟明显；不及时排粪类天牛幼虫运动能力相对较弱,表现为多无痛突,具细皱纹,侧纵沟浅。
天牛幼虫9对气门均呈长椭圆形或卵形。其中,中胸气门面积最大,其次是腹部末端气门面积,均显著大于其它7对气门面积比例值。排粪类天牛各体节气门面积占侧面积比例值显著大于不排粪类幼虫。
2、天牛内部形态特征与取食习性的关系
通过比较分析9种老熟天牛幼虫消化道特征与其取食树种的关系,以及呼吸系统特点与其排粪习性的关系；测量4种老熟幼虫呼吸速率与其排粪特征的关系。
明确天牛幼虫消化道结构简单,消化道长度显著长于体长。其中,中肠最长,占整个消化道长度的0.63-0.73；后肠次之,占0.22-0.35；前肠最短,不足消化道长度的0.06。危害针叶树的幼虫前肠占消化道总长比例均低于危害阔叶树的幼虫前肠比例。
9种天牛老熟幼虫呼吸系统结构特点为,体内有两条主气管,分布于两侧。气管的前端连接前胸,后端延伸至腹部最末端。气管与两侧气门相连但后胸气门不发达,仅具痕迹。气管与气门相连之处分出的二级、三级气管及微气管,与复眼、触角、消化道等器官相连。排粪类天牛幼虫一级气管直径大于不排粪幼虫直径。排粪类天牛幼虫二级气管数量较多,且直径大于不排粪类幼虫。
4种老熟幼虫,排粪类的桑天牛Apriona germari和松墨天牛Monochamus alternatus的幼虫代谢速率显著大于不排粪类的双条杉天牛Semanotus bifasciatus和红缘天牛Asias halodendri幼虫。上述4种天牛幼虫的呼吸墒值差异不大,但排粪幼虫呼吸墒值还是大于不排粪类幼虫。
总之,上述结果表明,天牛幼虫的形态结构与其取食习性存在明显的适应关系。
Cerambycinae which are important forest boring beetles, caused enormous losses on China's forestry by damaging plant tissues. A lot of scholars from home and abroad have done much work in Cerambycinae's characteristics, life history, life habits, spatial distribution pattern and digestive enzymes, but few people investigated the adaptability of morphological structures and harming habits of cerambycid larvae. The study observed the cerambycid larvae's morphological structures and the relationship with feeding habits, which included the main feeding site, the material of host plants and timely discharged faeces and sawdusts or not. The main results are as follow:
1、The relationship of external morphological structures and feeding habits:observed the mandible of15cerambycid species belong to3subfamilies of Cerambycidae and investigated the relationship of9mature larvae's developed degree of mandible and the material of host plants or the main feeding site; compared the relationship of the characteristics of the moving organs of16cerambycid larvae and discharging habits, and analyzed the relationship of9mature larvae's spiracles and discharging habits.
The results showed that the7larvae of subfamily Cerambyciae had gouge-shaped mandible, and the7larvae of subfamily Lamiinae and the1larvae of subfamily Aseminae had wedge-shaped mandible. From the morphology of the mandible's top, the back and inner surface of gouge-shaped mandible had several transverse furrows; the back of wedge-shaped mandible had several transverse furrows and its'inner surface had strong oncus. In the9species of mature larvae, For the larvae with gouge-shaped mandible, the smaller of the area ratio of its exerted-visible mandible to head, the stronger its mandible is. For the larvae with wedge-shaped mandible, the larger of the area ratio of its exerted-visible mandible to head, the stronger its mandible is. The mandible of larvae feeding broadleaf were stronger than the mandible of larvae feeding conifer, and the mandible of larvae mainly harming xylem were stronger than the mandible of larvae mainly harming phloem.
Observation of16species of larvae, the alar lobe of pronotum was strong, and had obvious lateral furrow, epipleurun and the first half of median line. Most of the larvae of subfamily Lamiinae had postnotum fold, the larvae of subfamily of Cerambyciae and Aseminae did not. The latter half of pronotum in the larvae which timely discharged faeces and sawdusts had'convex' pattern and tough particles or punctum, and which didn't timely discharge faeces and sawdusts had fine wrinkles and particles or small round spot. The ambulatory ampullae in the larvae which timely discharged faeces and sawdusts were more developed, with tough and big verrucae. The sports ability in the larvae which did not timely discharged faeces and sawdusts were worse, with fine wrinkle and small verrucae.
All the spiracles of cerambycid larvae were long oval or oval. Among them, the relative area of mesostigma was the largest, followed by the area of abdominal end spiracles, were significantly greater than the other7on the spiracles area ratio.the relative area of spiracles in the larvae which timely discharged faeces and sawdusts were larger than the larvae which did not.
2、The relationship of the inside morphological structures and feeding habits:observed the characteristic of alimentary canal of9mature cerambycid larvae and the relationship of the material of host plants, investigated the characteristic of the respiratory systems of9mature cerambycid larvae and the relationship of discharging habits, measured the respiration rate of4mature cerambycid larvae and the relationship of discharging faeces and sawdusts.
The structure of larvae's alimentary canal was simple, and its'length was obviously longer than the body. Among them, the midgut was the longest, accouting for the alimentary canal length0.63-0.73; hindgut followed by accounting for0.22-0.35; foregut was the shortest, less than0.06of the whole length. For the larvae feeding conifer, its'ratio of foregut to alimentary canal was smaller than the larvae feeding broadleaf.
The characteristic of the respiratory systems of9mature larvae; the body had two main trachea, distributed in both sides. The front of trachea connected to the prothorax, and the back-end extended to the end of the abdominal. The trachea connected spiracles on both sides, but metastigma was not developed, with only trace. The connection point of trachea and spiracles separated the secondary trachea, thirdary trachea and tracheole, and connected to compound eyes, antenna, alimentary canal and other organs. The primary trachea's diameter of larvae timely discharged faeces and sawdusts was larger than the larvae did not. The larvae timely discharged faeces and sawdusts had a larger number of secondary trachea, and its'diameter were larger than the larvae did not timely discharged.
In the4mature larvae, the larvae which timely discharged faeces and sawdusts-the metabolic rate of Apriona germari and Monochamus alternates were larger than the larvae did not timely discharged-Semanotus bifasciatus and Asias halodendri. The respiratory entropy of4species of cerambycid larvae were not significantly different, but respiratory entropy of larvae timely discharged was larger than the larvae did not timely discharged.
In all, the results above showed that the cerambycid larvae's morphological structures and feeding habits obviously had adaptive relationship.
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