中国锯天牛亚科比较形态学研究
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摘要
锯天牛亚科(Prioninae)隶属于鞘翅目(Coleoptera)天牛总科(Cerambycoidea)天牛科(Cerambycidae),不少种类是农林业中的重要害虫,具有十分重要的经济意义。本论文介绍了锯天牛亚科的经济意义和国内外研究现状,天牛总科后翅、雌雄性生殖器的研究进展,比较形态学研究现状等内容：本文针对我国锯天牛亚科研究的不足以及锯天牛亚科分类系统的不统一,比较研究了锯天牛亚科的外部形态特征、后翅翅脉特征、雌雄性生殖器特征和雄性第8节腹板形态特征：筛选出了锯天牛亚科在族级、属级和种级具有分类学意义的特征；主要研究结果如下：
1传统外部形态特征比较研究
本研究分别从头胸腹选取了触角的长短、前胸背板侧齿的数量、腹部末节端部的形状等94个形态特征,比较研究了中国锯天牛亚科5族14属47种和亚种188号昆虫标本。发现了1个新纪录种：Anomophysis ellioti (Waterhouse,1884)艾略特异胸天牛。通过比较研究,筛选出了外部形态具有分类学意义的特征,如触角的形状、触角的节数、鞘翅内端角的形状和前胸背部侧齿的数量等68个特征；同时提出了避免使用鞘翅颜色(差异较小)等特征做为分类特征的建议。拍摄了20张特征形态图。
2后翅翅脉特征比较研究
本研究选取了径室Rc的形状等后翅翅脉特征22个,详细比较研究了中国锯天牛亚科5族14属47种和亚种75号后翅标本。通过对后翅翅脉特征的比较研究发现,后翅翅脉特征在族级阶元具有明显的分类学意义,在属级和种级分类意义不明显。筛选出了锯天牛亚科具有分类意义的后翅翅脉特征,如径室Rc的形状、楔状室的形状和CuA1+2脉是否存在等7个特征。拍摄及处理了中国锯天牛亚科5族14属47种和亚种75号后翅翅脉图片。
3雌雄性生殖器特征比较研究
本文选取了阳基侧突端部形状等26个雄性外生殖器特征,解剖比较研究了中国锯天牛亚科5族11属36种和亚种38号雄性外生殖器标本：选取了基腹片突形状等20个雌性生殖器特征,解剖研究了5族11属22种和亚种22号雌性生殖器标本。通过比较研究发现,锯天牛亚科雌雄性外生殖器特征在族级、属级和种(亚种)级均具有分类学意义;同时,雌雄性生殖器特征还可以用于近似种的区分。筛选出了雌雄性生殖器具有分类学意义的特征(雌性生殖器包括载肛突骨杆的有无和阴道片的长短等16个特征,雄性外生殖器包括阳基侧突端部的形状和盖的有无等24个特征)。拍摄并处理了中国锯天牛亚科5族11属36种(亚种)38号雄性外生殖器阳基侧突、环端部和中茎照片,5族11属22种(亚种)22号雌性生殖器产卵器的背面、腹面和内生殖器照片。
4雄性第8节腹板比较研究
本文首次对锯天牛亚科雄性第8节腹板的形态特征进行了详细的比较研究。选取了6个特征对4族9属31种和亚种33号标本进行了比较研究。研究发现锯天牛亚科雄性第8节腹板在种间具有明显的差异性,但它们不具相同的属征和族征；因此,在属级及族级不具分类学意义。拍摄并处理了中国锯天牛亚科4族9属31种(亚种)33号雄性第8节腹板的形态图片。
5比较形态学研究结果
通过比较形态学研究,筛选出了锯天牛亚科在族级、属级和种级具有分类学意义的特征。族级具有分类学意义的特征主要有：后胸前侧片两侧近于平行或显著向后狭窄,触角的形状(扁形或圆柱形),前胸背板侧齿的数量,腹部最后1节可见腹板端部的形状,产卵管是否外露。后翅翅脉具有分类意义的特征：径室的形状,RP脉的长短(RP脉/r4脉),MP1+2脉端半部的长短(r-m/MP1+2端半部长度),CuA1+2脉是否存在等。雄性外生殖器具有分类意义的特征：盖的有无及长短,环向腹面弯折的角度大小,交配孔腹缘是否突出等。雌性生殖器具有分类意义的特征：产卵器的骨化程度(背骨杆是否骨化呈骨板),载肛突骨杆的长短,基腹片突的大小及形状,端突的着生位置等。
属级具有分类学意义的特征主要有：触角的节数,触角第3节腹面是否有凹槽,上颚是否细长,上颚基部是否被毛,头表面是否被毛,前胸背板是否具侧齿,小盾片是否被毛,第3跗节的分裂情况(正常,很浅,很深),腹部是否光滑等。雄性外生殖器阳基的长短(与中茎比较),阳基侧突端部的形状,阳基侧突刚毛的疏密,环在端部是否合并,环向腹面弯折的位置,中茎基的长短(与中茎突比较)等。受精囊腺在受精囊上的着生位置,阴道片的长短及骨化程度,基腹片骨杆和负瓣片骨杆是否分离,背骨杆的有无等。
种级具有分类学意义的特征主要有：上颚基部是否被毛及外缘是否具齿突,下唇须和下颚须末节的形状,触角的节数及长短,前胸腹板突片是否显著突出于基节之上,前胸背板是否具侧齿及侧齿的数量,鞘翅表面是否被毛,鞘翅肩角是否具齿,中后足胫节背面是否具棱脊,第3跗节的形状等。雄性外生殖器阳基侧突端部的形状,阳基侧突刚毛的长短及疏密,环向腹面的弯折位置,环端部合并处是否收狭,中茎是否向腹面弯曲,中茎基端部的形状,中茎突端部的形状,交配孔背缘中部的形状等特征。雌性生殖器受精囊的形状,交配囊的形状,肛侧板骨杆的长短(肛侧板骨杆/产卵器),背骨杆的长短(背骨杆/产卵器),背骨杆基部是否有分支,载肛突骨杆的有无等。雄性第8节腹板在种间也具有差异性,主要体现在端部的形状,长宽比例,端部是否收狭等特征上。
依据比较形态学研究结果和参考前人资料,结合目前最新的分类研究观点,编制了中国锯天牛亚科6族23属105种(亚种)的分类检索表。本研究不仅为中国锯天牛亚科昆虫的分类鉴定提供了更准确、可靠的分类特征,还为其系统发育及进化等深入研究奠定了一定的理论基础。
Prioninae belongs to Cerambycidae of Cerambycoidea in Coleoptera, and some species from this subfamily are important insect pests in agriculture and forest, having important economic significance. This thesis introduces the economic significance and the research progresses in Prioninae, recent developments of hind-wing venations, productive organs and comparative morphology study. However, the research in Prioninae is insufficient, and some researchers declared different viewpoint in the classification system of Prioninae. Therefore, in the current research, comparative morphology study was carried out in Prioninae focusing on the external morphology, hind-wing venations, productive organs and the 8th sternum. Many characters were screen out in tribes, genera and species of Prioninae. The main results are following:
1 Comparative study on the traditional external morphology of Prioninae
In the current study,94 characters of traditional external morphology were chose to make the comparative research on 188 specimens of 47 species and subspecies of 14 genera in 5 tribes. Anomophysis ellioti (Waterhouse,1884) was recorded at the first time in China.68 characters of classification were also found through comparative research and some confused characters should be avoided in the further research. The pictures of 20 characters were presented.
2 Comparative study on the hind-wing venations of Prioninae
21 characters of hind-wing venations were chose to make the comparative study on 75 specimens of 47 species and subspecies of 14 genera in 5 tribes. The results revealed that the characters of hind-wing venation can be used to distinguish the tribes of Prioninae, but it is not obvious in the genus and species. In addition,75 pictures were obtained in 5 tribes,14 genera,47 species and subspecies.
3 Comparative study on the reproductive organs of Prioninae
In our research,26 characters of male genitalia were chose to make the comparative research on 38 specimens of 36 species and subspecies of 11 genera in 5 tribes, while 20 characters of female genital organs were chose to make the comparative research on 22 specimens of 22 species and subspecies of 11 genera in 5 tribes. The results indicated that the characters have the taxonomic significance in tribe, genus and species. All pictures were also obtained.
4 Comparative study on the 8th sternum of male of Prioninae
The 8th sternum of Prioninae was first used in our comparative study. Six characters of the 8th sternum were chose to make the comparative study on 33 speciemens of 31 species and subspecies of 9 genera of 4 tribes. The results suggested that the characters of the 8th sternum showed difference in species, but they do not have the same characters of genus. The pictures of the 8th sternum were also obtained.
5 The results of comparative morphology study
The taxonomic characters were selected in species, genus and tribe through the whole research. The important characters that can be used to distinguish the tribes are both sides of front lateral plate of metathorax nearly parallel or stricture, the shape of antenna, the numbers of latadentis, the shape of Re, the length of the RP, the roof exist or not, the length of proctiger baculum, the shape and size of the coxite lobe, and so on.
Many taxonomic characters were also found to be used in distinguishing genera. The main taxonomic characters as following:the numbers of segments of antenna, existing groove or not, the length of the tegmen, the density of the seta in lateral lobes of male genitalia, the size and the degree of sclerotization of the vaginal plate, having dorsal baculum or not in female genital organ, and so on.
In addition, some characters can be used to distinguish species and subspecies, such as the shape of the last segment of labipalp and maxillary palp, the numbers of segment of antenna, existing hair on the elytron or not, the shape of the 3th tarsi, the top shape of the lateral lobes, the shape of the spermatheca, the shape of the bursa copulatrix, the size of the dorsal baculum, having proctiger baculum or not, the shape of the apex of the 8th sternum, and so on.
Based on our research and the previous materials, an identification key to species and subspecies of Prioninae from China is presented. This thesis established the theoretical basis for intensive studv of Prioninae in China.
1传统外部形态特征比较研究
本研究分别从头胸腹选取了触角的长短、前胸背板侧齿的数量、腹部末节端部的形状等94个形态特征,比较研究了中国锯天牛亚科5族14属47种和亚种188号昆虫标本。发现了1个新纪录种：Anomophysis ellioti (Waterhouse,1884)艾略特异胸天牛。通过比较研究,筛选出了外部形态具有分类学意义的特征,如触角的形状、触角的节数、鞘翅内端角的形状和前胸背部侧齿的数量等68个特征；同时提出了避免使用鞘翅颜色(差异较小)等特征做为分类特征的建议。拍摄了20张特征形态图。
2后翅翅脉特征比较研究
本研究选取了径室Rc的形状等后翅翅脉特征22个,详细比较研究了中国锯天牛亚科5族14属47种和亚种75号后翅标本。通过对后翅翅脉特征的比较研究发现,后翅翅脉特征在族级阶元具有明显的分类学意义,在属级和种级分类意义不明显。筛选出了锯天牛亚科具有分类意义的后翅翅脉特征,如径室Rc的形状、楔状室的形状和CuA1+2脉是否存在等7个特征。拍摄及处理了中国锯天牛亚科5族14属47种和亚种75号后翅翅脉图片。
3雌雄性生殖器特征比较研究
本文选取了阳基侧突端部形状等26个雄性外生殖器特征,解剖比较研究了中国锯天牛亚科5族11属36种和亚种38号雄性外生殖器标本：选取了基腹片突形状等20个雌性生殖器特征,解剖研究了5族11属22种和亚种22号雌性生殖器标本。通过比较研究发现,锯天牛亚科雌雄性外生殖器特征在族级、属级和种(亚种)级均具有分类学意义;同时,雌雄性生殖器特征还可以用于近似种的区分。筛选出了雌雄性生殖器具有分类学意义的特征(雌性生殖器包括载肛突骨杆的有无和阴道片的长短等16个特征,雄性外生殖器包括阳基侧突端部的形状和盖的有无等24个特征)。拍摄并处理了中国锯天牛亚科5族11属36种(亚种)38号雄性外生殖器阳基侧突、环端部和中茎照片,5族11属22种(亚种)22号雌性生殖器产卵器的背面、腹面和内生殖器照片。
4雄性第8节腹板比较研究
本文首次对锯天牛亚科雄性第8节腹板的形态特征进行了详细的比较研究。选取了6个特征对4族9属31种和亚种33号标本进行了比较研究。研究发现锯天牛亚科雄性第8节腹板在种间具有明显的差异性,但它们不具相同的属征和族征；因此,在属级及族级不具分类学意义。拍摄并处理了中国锯天牛亚科4族9属31种(亚种)33号雄性第8节腹板的形态图片。
5比较形态学研究结果
通过比较形态学研究,筛选出了锯天牛亚科在族级、属级和种级具有分类学意义的特征。族级具有分类学意义的特征主要有：后胸前侧片两侧近于平行或显著向后狭窄,触角的形状(扁形或圆柱形),前胸背板侧齿的数量,腹部最后1节可见腹板端部的形状,产卵管是否外露。后翅翅脉具有分类意义的特征：径室的形状,RP脉的长短(RP脉/r4脉),MP1+2脉端半部的长短(r-m/MP1+2端半部长度),CuA1+2脉是否存在等。雄性外生殖器具有分类意义的特征：盖的有无及长短,环向腹面弯折的角度大小,交配孔腹缘是否突出等。雌性生殖器具有分类意义的特征：产卵器的骨化程度(背骨杆是否骨化呈骨板),载肛突骨杆的长短,基腹片突的大小及形状,端突的着生位置等。
属级具有分类学意义的特征主要有：触角的节数,触角第3节腹面是否有凹槽,上颚是否细长,上颚基部是否被毛,头表面是否被毛,前胸背板是否具侧齿,小盾片是否被毛,第3跗节的分裂情况(正常,很浅,很深),腹部是否光滑等。雄性外生殖器阳基的长短(与中茎比较),阳基侧突端部的形状,阳基侧突刚毛的疏密,环在端部是否合并,环向腹面弯折的位置,中茎基的长短(与中茎突比较)等。受精囊腺在受精囊上的着生位置,阴道片的长短及骨化程度,基腹片骨杆和负瓣片骨杆是否分离,背骨杆的有无等。
种级具有分类学意义的特征主要有：上颚基部是否被毛及外缘是否具齿突,下唇须和下颚须末节的形状,触角的节数及长短,前胸腹板突片是否显著突出于基节之上,前胸背板是否具侧齿及侧齿的数量,鞘翅表面是否被毛,鞘翅肩角是否具齿,中后足胫节背面是否具棱脊,第3跗节的形状等。雄性外生殖器阳基侧突端部的形状,阳基侧突刚毛的长短及疏密,环向腹面的弯折位置,环端部合并处是否收狭,中茎是否向腹面弯曲,中茎基端部的形状,中茎突端部的形状,交配孔背缘中部的形状等特征。雌性生殖器受精囊的形状,交配囊的形状,肛侧板骨杆的长短(肛侧板骨杆/产卵器),背骨杆的长短(背骨杆/产卵器),背骨杆基部是否有分支,载肛突骨杆的有无等。雄性第8节腹板在种间也具有差异性,主要体现在端部的形状,长宽比例,端部是否收狭等特征上。
依据比较形态学研究结果和参考前人资料,结合目前最新的分类研究观点,编制了中国锯天牛亚科6族23属105种(亚种)的分类检索表。本研究不仅为中国锯天牛亚科昆虫的分类鉴定提供了更准确、可靠的分类特征,还为其系统发育及进化等深入研究奠定了一定的理论基础。
Prioninae belongs to Cerambycidae of Cerambycoidea in Coleoptera, and some species from this subfamily are important insect pests in agriculture and forest, having important economic significance. This thesis introduces the economic significance and the research progresses in Prioninae, recent developments of hind-wing venations, productive organs and comparative morphology study. However, the research in Prioninae is insufficient, and some researchers declared different viewpoint in the classification system of Prioninae. Therefore, in the current research, comparative morphology study was carried out in Prioninae focusing on the external morphology, hind-wing venations, productive organs and the 8th sternum. Many characters were screen out in tribes, genera and species of Prioninae. The main results are following:
1 Comparative study on the traditional external morphology of Prioninae
In the current study,94 characters of traditional external morphology were chose to make the comparative research on 188 specimens of 47 species and subspecies of 14 genera in 5 tribes. Anomophysis ellioti (Waterhouse,1884) was recorded at the first time in China.68 characters of classification were also found through comparative research and some confused characters should be avoided in the further research. The pictures of 20 characters were presented.
2 Comparative study on the hind-wing venations of Prioninae
21 characters of hind-wing venations were chose to make the comparative study on 75 specimens of 47 species and subspecies of 14 genera in 5 tribes. The results revealed that the characters of hind-wing venation can be used to distinguish the tribes of Prioninae, but it is not obvious in the genus and species. In addition,75 pictures were obtained in 5 tribes,14 genera,47 species and subspecies.
3 Comparative study on the reproductive organs of Prioninae
In our research,26 characters of male genitalia were chose to make the comparative research on 38 specimens of 36 species and subspecies of 11 genera in 5 tribes, while 20 characters of female genital organs were chose to make the comparative research on 22 specimens of 22 species and subspecies of 11 genera in 5 tribes. The results indicated that the characters have the taxonomic significance in tribe, genus and species. All pictures were also obtained.
4 Comparative study on the 8th sternum of male of Prioninae
The 8th sternum of Prioninae was first used in our comparative study. Six characters of the 8th sternum were chose to make the comparative study on 33 speciemens of 31 species and subspecies of 9 genera of 4 tribes. The results suggested that the characters of the 8th sternum showed difference in species, but they do not have the same characters of genus. The pictures of the 8th sternum were also obtained.
5 The results of comparative morphology study
The taxonomic characters were selected in species, genus and tribe through the whole research. The important characters that can be used to distinguish the tribes are both sides of front lateral plate of metathorax nearly parallel or stricture, the shape of antenna, the numbers of latadentis, the shape of Re, the length of the RP, the roof exist or not, the length of proctiger baculum, the shape and size of the coxite lobe, and so on.
Many taxonomic characters were also found to be used in distinguishing genera. The main taxonomic characters as following:the numbers of segments of antenna, existing groove or not, the length of the tegmen, the density of the seta in lateral lobes of male genitalia, the size and the degree of sclerotization of the vaginal plate, having dorsal baculum or not in female genital organ, and so on.
In addition, some characters can be used to distinguish species and subspecies, such as the shape of the last segment of labipalp and maxillary palp, the numbers of segment of antenna, existing hair on the elytron or not, the shape of the 3th tarsi, the top shape of the lateral lobes, the shape of the spermatheca, the shape of the bursa copulatrix, the size of the dorsal baculum, having proctiger baculum or not, the shape of the apex of the 8th sternum, and so on.
Based on our research and the previous materials, an identification key to species and subspecies of Prioninae from China is presented. This thesis established the theoretical basis for intensive studv of Prioninae in China.
引文
[1]陈力,李竹,冯波.中国锯天牛亚科区系研究.西南大学学报(自然科学版),2010,32(4)：1-5.
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