中国锹甲科部分属种鞘翅和雄性外生殖器外翻囊的超微形态研究（鞘翅目：金龟总科）
摘要
锹甲是鞘翅目Coleoptera金龟总科Scarabaeoidea中较为原始的类群。绝大多数种类都以森林为栖息地。幼虫多取食分解朽木,在生态系统的物质能量循环中占据独特的生态位。是一类相当值得研究的昆虫。但作为各项研究基础的分类学研究总体较弱,存在很多问题。特别是同一种内所具有的雌雄二型、雄性多型及不同的色斑型,给物种鉴定带来很大困难,出现了一系列疑难属种。鉴于此,本文尝试利用扫描电镜技术,比较了中国产4亚科15属75种(含亚种)锹甲的鞘翅与雄性外生殖器外翻囊的超微形态,以探索鞘翅与雄性外生殖器外翻囊的超微形态在种类鉴定及探讨高级阶元系统关系上意义。相关研究结果如下：
在亚科阶元上,通过比较鞘翅表面毛窝伴生孔的有无,可以发现：
1.锹甲亚科Lucaninae的代表性属Lucanus鞘翅表面毛窝无伴生孔,而刀锹甲亚科的代表性属如广义刀锹甲属Dorcus (s.s)中的4属鞘翅表面毛窝均具伴生孔,说明刀锹甲亚科Dorcinae不能归入锹甲亚科,即不支持Holloway(2007)的观点。
2.盾锹甲属Aegus、柱锹甲属Prismognathus、环锹甲属Cyclommatus的毛窝均无伴生孔,说明也不能归入刀锹甲亚科即不支持Bartolozzi&Sprecher (2006)的观点。
3.部分前锹甲属Prosopocoilus的种(如儒圣前锹甲P. confucius、缝前锹甲P.suturalis、双斑前锹甲P. biplagiatus、歧齿前锹甲P. approximates、剪齿前锹甲P.forficulaforficula、黄褐前锹甲P. blanchardi)的毛窝也无伴生孔,毛窝呈光滑的、深椭圆形毛孔状,刚毛着生在一侧,说明这些种也不能归入刀锹甲亚科。4.奥锹甲亚科Odontolabinae(新锹甲属Neolucanus、奥锹甲属Odontolabis)的鞘翅表面毛窝无伴生孔,毛窝呈深椭圆形的毛孔状,紧邻毛窝一侧具密集的颗粒状突起及褶皱状脊纹,刚毛曲向一侧。这些特征说明奥锹甲亚科确实应为一个独立业科,不能归入刀锹甲亚科和锹甲亚科,即不支持Holloway (2007)的观点。
在属级阶元上,比较毛窝的结构形状及其与伴生孔的距离,可得出以下结论：
1.广义刀锹甲属Dorcus (s.l.)中的4属所研究的种类鞘翅毛窝均具有伴生孔,伴生孔距离毛窝较远,说明它们的确是近缘属,其中：1)莫锹甲属Macrodorcas不应作为一个独立有效的属：该属的模式种M. recta和狭义刀锹甲属Dorcus (s.s.)的典型种类D. davidis具几乎完全相同的毛窝结构(即毛窝都呈近深凹的圆形,密布楞脊状突起)。这表明其不应当作为一个独立有效的属。这与我们之前的部分系统发育研究结果相一致。2)半刀锹甲属Hemisodorcus应作为一个独立有效的属：本研究中的半刀锹甲属Hemisodorcus的模式种H. nepalensis及其它种的鞘翅毛窝都呈较细而深的毛孔状,相当光滑,明显不同于狭义刀锹甲属Dorcus (s. s.)和扁锹甲属Serrognathus的种类,表明该属应作为一个独立有效属。莫锹甲属Macrodorcas中其他种如M. pieli、M. melliana、M. seguyi、M. mochizukii、M. elsiedis及部分前锹甲属Prosopocoilus的种如P. gracilis、P. denticulutus、P. crenulimens、P. fomosanus、P. melli的鞘翅毛窝都呈细深的毛孔状且相当光滑,表明它们可归入半刀锹甲属中。3)锈色刀锹甲D. velutinus的系统地位需重新讨论：该种的鞘翅毛窝呈两种类型,鞘翅表面具针状突,刚毛羽扇状。这些超微形态特征为D. velutinus所特有,且明显不同于本文研究的其它种。因此,这个种的系统地位需重新讨论。
2.六节锹甲属Hexarthrius、拟鹿锹甲属Pseudorhaetus、鹿角锹甲属Rhaetulus及部分前锹甲属Prosopocoilus的种类(如刺前锹甲P. spineus、阿斯前锹甲P. astacoides)的鞘翅毛窝均具伴生孔,且孔距毛窝较近。这表明它们亲缘关系较近。
3.前锹甲属Prosopocoilus的分类需要重新厘定。如前所述,本文研究的中国产前锹甲属的种类可以分为3大类,即鞘翅不具伴生孔、鞘翅毛窝距伴生孔较远、鞘翅毛窝距伴生孔较近等。因而,需要对这个属目前包括的种重新进行分类厘定。
在种级水平上,鞘翅表面有无龟裂块及显著程度、刚毛的长短与形状及雄外生殖器外翻囊的超微形态适于种间差异研究。综合比较这些特征,可得出以下结论：基锐前锹甲P. denticulatus与基叉前锹甲P. crenulidens应分别为两个独立有效种；三齿新锹甲N. delicatus在同一种内确实存在两个色斑型；中华奥锹甲O.cuvera sinensis和华美奥锹甲O. cuvera fallaciosus可以作为两个独立种。
Stag beetles belong to one of primitive families in Scarabaeoidea (Coleoptera). Most of them habituated in forest. Larvae of them live in rotten logs and fed off the humus so that they occupied very special niche of linking the energy flow among the ecosystem, which indicates they are worth researching. However, as the foundational work, it is not so clear about the taxonomy of stag beetles in the world. Especially. a lot of problematic genera and species need to be clarified, which usually caused by intraspecific traits such as sexual dimorphism and male polymorphism, or highly interspecific similarity. In this thesis, SEM was used to obtain the ultrastructure of elytra and PES of stag beetles. Our aim is to search some useful characters for identifying problematic species and revealing the relationships among higher taxa in Lucanidae based on studying four subfamilies, fifteen genera and75species (including subspecies) from China. The conclusions are presented as follows:
On the level of subfamilies:1. Dorcinae shall be a valid subfamily and cannot be included in Lucaninae because of elytra setal pits with accompanying holes in Dorcinae; whereas, elytra setal pits in Lucaninae without accompanying holes, i.e., the conclusion does not support the opinion of Holloway in2007.2. The three genera, Aegus, Prismognathus and Cyclommatus cannot be taken into Dorcinae due to accompanying hole are absent on the elytra, i.e., the conclusion does not support the opinion of Bartolozzi&Sprecher in2006.3. Partly members of Prosopocoilus, such as, P. confucius, P. suturalis, P. biplagiatus, P. approximates, P. forficula forfiaila, P. blanchardi, cannot be in Dorcinae considering their accompanying holes are absent and setal pits are different.4. Odontolabinae which formed by Neolucanus and Odontolabis cannot be included in Lucaninae, too. Elytra setal pits of all the studied species from the two genera present deeply oblong pore-shape with densely granular produces and wrinkly folds which closed to the side where the elytral setae bent to. The accompanying holes are absent. These characters support that Odontolabinae actually is a valid subfamily because of these special characters. It cannot be moved into Lucaninae as Holloway had claimed in2007.
On the level of genera:1. Elytra of four members of Dorcus (s.l.) share the same character, i.e., each setal pit process a distinctly distant accompanying hole. The following conclusions can be presented about them:1) The genus Macrodorcas could be invalid status because the setal pits of its generic type(M. recta) are almost same as that of Dorcus davidis which is a typical species of Dorcus (s.s). The conclusion is in accord with our known phylogeny analysis in2012.2) Hemisodorcus could be a valid genus because the setal pits of its generic type species (H. nepalensis) are deeply pore-shape, very smooth, which is distinctly different from those of Dorcus (s. s.) and Serrognathus. Nevertheless, the character is almost same as those of some species in Macrodorcas (M pieli, M. melliana, M. seguyi, M. mochizukii, M. elsiedis) and some species in Prosopocoilus(P. gracilis, P. denticulutus, P. cremilimens, P. fomosanus, P. melli). These species could be members of Hemisodorcus.3) Taxonomy of Dorcus velutinus need discussed re-newly because of two kinds of setal pit, densely needlelike tips and fan-shaped setae on the elytra surface. These characters indicated Dorcus velutinus could has new status.2. Each setal pit is immediately to each accompanying hole in Hexarthrius, Pseudorhaetus, Rhaetulus and some species of Prosopocoilus(P. spineus, P. astacoides), which can tell that they have more closer relationships.3. Taxonomy of the present Prosopocoilus shall be clarified re-newly because the above-mentioned result. All the studied species in this genus from China can be separated into three groups-based on setal pits characters.
On the species level:the following characters on the elytra, flattened polygonal mounds present or absent, marked degree (easy to recognize or dim shape), setae length and shape, are useful to differ species. Also, the length, shape, distribution and adherent direction of affiliated hairs on the apical part of PES exhibited apparently differences among species. Some problematic species can be differed distinctly by analyzing the combination these elytra with PES ultrastructure:P. denticulatus is not a synonym of P. crenulidens as Benesh wrote in1960, but two valid species due to their remarkably different affiliated hairs; elytra and PES ultrastructure between yellow spot and whole dark elytra individuals of N. delicatus showed very highly similarity, which confirmed that two different splash types are actually existed; O. cuvera sinensis and O. cuvera fallaciosus could be two good species because the shapes and adherent direction between their affiliated hairs are conspicuously different.
在亚科阶元上,通过比较鞘翅表面毛窝伴生孔的有无,可以发现：
1.锹甲亚科Lucaninae的代表性属Lucanus鞘翅表面毛窝无伴生孔,而刀锹甲亚科的代表性属如广义刀锹甲属Dorcus (s.s)中的4属鞘翅表面毛窝均具伴生孔,说明刀锹甲亚科Dorcinae不能归入锹甲亚科,即不支持Holloway(2007)的观点。
2.盾锹甲属Aegus、柱锹甲属Prismognathus、环锹甲属Cyclommatus的毛窝均无伴生孔,说明也不能归入刀锹甲亚科即不支持Bartolozzi&Sprecher (2006)的观点。
3.部分前锹甲属Prosopocoilus的种(如儒圣前锹甲P. confucius、缝前锹甲P.suturalis、双斑前锹甲P. biplagiatus、歧齿前锹甲P. approximates、剪齿前锹甲P.forficulaforficula、黄褐前锹甲P. blanchardi)的毛窝也无伴生孔,毛窝呈光滑的、深椭圆形毛孔状,刚毛着生在一侧,说明这些种也不能归入刀锹甲亚科。4.奥锹甲亚科Odontolabinae(新锹甲属Neolucanus、奥锹甲属Odontolabis)的鞘翅表面毛窝无伴生孔,毛窝呈深椭圆形的毛孔状,紧邻毛窝一侧具密集的颗粒状突起及褶皱状脊纹,刚毛曲向一侧。这些特征说明奥锹甲亚科确实应为一个独立业科,不能归入刀锹甲亚科和锹甲亚科,即不支持Holloway (2007)的观点。
在属级阶元上,比较毛窝的结构形状及其与伴生孔的距离,可得出以下结论：
1.广义刀锹甲属Dorcus (s.l.)中的4属所研究的种类鞘翅毛窝均具有伴生孔,伴生孔距离毛窝较远,说明它们的确是近缘属,其中：1)莫锹甲属Macrodorcas不应作为一个独立有效的属：该属的模式种M. recta和狭义刀锹甲属Dorcus (s.s.)的典型种类D. davidis具几乎完全相同的毛窝结构(即毛窝都呈近深凹的圆形,密布楞脊状突起)。这表明其不应当作为一个独立有效的属。这与我们之前的部分系统发育研究结果相一致。2)半刀锹甲属Hemisodorcus应作为一个独立有效的属：本研究中的半刀锹甲属Hemisodorcus的模式种H. nepalensis及其它种的鞘翅毛窝都呈较细而深的毛孔状,相当光滑,明显不同于狭义刀锹甲属Dorcus (s. s.)和扁锹甲属Serrognathus的种类,表明该属应作为一个独立有效属。莫锹甲属Macrodorcas中其他种如M. pieli、M. melliana、M. seguyi、M. mochizukii、M. elsiedis及部分前锹甲属Prosopocoilus的种如P. gracilis、P. denticulutus、P. crenulimens、P. fomosanus、P. melli的鞘翅毛窝都呈细深的毛孔状且相当光滑,表明它们可归入半刀锹甲属中。3)锈色刀锹甲D. velutinus的系统地位需重新讨论：该种的鞘翅毛窝呈两种类型,鞘翅表面具针状突,刚毛羽扇状。这些超微形态特征为D. velutinus所特有,且明显不同于本文研究的其它种。因此,这个种的系统地位需重新讨论。
2.六节锹甲属Hexarthrius、拟鹿锹甲属Pseudorhaetus、鹿角锹甲属Rhaetulus及部分前锹甲属Prosopocoilus的种类(如刺前锹甲P. spineus、阿斯前锹甲P. astacoides)的鞘翅毛窝均具伴生孔,且孔距毛窝较近。这表明它们亲缘关系较近。
3.前锹甲属Prosopocoilus的分类需要重新厘定。如前所述,本文研究的中国产前锹甲属的种类可以分为3大类,即鞘翅不具伴生孔、鞘翅毛窝距伴生孔较远、鞘翅毛窝距伴生孔较近等。因而,需要对这个属目前包括的种重新进行分类厘定。
在种级水平上,鞘翅表面有无龟裂块及显著程度、刚毛的长短与形状及雄外生殖器外翻囊的超微形态适于种间差异研究。综合比较这些特征,可得出以下结论：基锐前锹甲P. denticulatus与基叉前锹甲P. crenulidens应分别为两个独立有效种；三齿新锹甲N. delicatus在同一种内确实存在两个色斑型；中华奥锹甲O.cuvera sinensis和华美奥锹甲O. cuvera fallaciosus可以作为两个独立种。
Stag beetles belong to one of primitive families in Scarabaeoidea (Coleoptera). Most of them habituated in forest. Larvae of them live in rotten logs and fed off the humus so that they occupied very special niche of linking the energy flow among the ecosystem, which indicates they are worth researching. However, as the foundational work, it is not so clear about the taxonomy of stag beetles in the world. Especially. a lot of problematic genera and species need to be clarified, which usually caused by intraspecific traits such as sexual dimorphism and male polymorphism, or highly interspecific similarity. In this thesis, SEM was used to obtain the ultrastructure of elytra and PES of stag beetles. Our aim is to search some useful characters for identifying problematic species and revealing the relationships among higher taxa in Lucanidae based on studying four subfamilies, fifteen genera and75species (including subspecies) from China. The conclusions are presented as follows:
On the level of subfamilies:1. Dorcinae shall be a valid subfamily and cannot be included in Lucaninae because of elytra setal pits with accompanying holes in Dorcinae; whereas, elytra setal pits in Lucaninae without accompanying holes, i.e., the conclusion does not support the opinion of Holloway in2007.2. The three genera, Aegus, Prismognathus and Cyclommatus cannot be taken into Dorcinae due to accompanying hole are absent on the elytra, i.e., the conclusion does not support the opinion of Bartolozzi&Sprecher in2006.3. Partly members of Prosopocoilus, such as, P. confucius, P. suturalis, P. biplagiatus, P. approximates, P. forficula forfiaila, P. blanchardi, cannot be in Dorcinae considering their accompanying holes are absent and setal pits are different.4. Odontolabinae which formed by Neolucanus and Odontolabis cannot be included in Lucaninae, too. Elytra setal pits of all the studied species from the two genera present deeply oblong pore-shape with densely granular produces and wrinkly folds which closed to the side where the elytral setae bent to. The accompanying holes are absent. These characters support that Odontolabinae actually is a valid subfamily because of these special characters. It cannot be moved into Lucaninae as Holloway had claimed in2007.
On the level of genera:1. Elytra of four members of Dorcus (s.l.) share the same character, i.e., each setal pit process a distinctly distant accompanying hole. The following conclusions can be presented about them:1) The genus Macrodorcas could be invalid status because the setal pits of its generic type(M. recta) are almost same as that of Dorcus davidis which is a typical species of Dorcus (s.s). The conclusion is in accord with our known phylogeny analysis in2012.2) Hemisodorcus could be a valid genus because the setal pits of its generic type species (H. nepalensis) are deeply pore-shape, very smooth, which is distinctly different from those of Dorcus (s. s.) and Serrognathus. Nevertheless, the character is almost same as those of some species in Macrodorcas (M pieli, M. melliana, M. seguyi, M. mochizukii, M. elsiedis) and some species in Prosopocoilus(P. gracilis, P. denticulutus, P. cremilimens, P. fomosanus, P. melli). These species could be members of Hemisodorcus.3) Taxonomy of Dorcus velutinus need discussed re-newly because of two kinds of setal pit, densely needlelike tips and fan-shaped setae on the elytra surface. These characters indicated Dorcus velutinus could has new status.2. Each setal pit is immediately to each accompanying hole in Hexarthrius, Pseudorhaetus, Rhaetulus and some species of Prosopocoilus(P. spineus, P. astacoides), which can tell that they have more closer relationships.3. Taxonomy of the present Prosopocoilus shall be clarified re-newly because the above-mentioned result. All the studied species in this genus from China can be separated into three groups-based on setal pits characters.
On the species level:the following characters on the elytra, flattened polygonal mounds present or absent, marked degree (easy to recognize or dim shape), setae length and shape, are useful to differ species. Also, the length, shape, distribution and adherent direction of affiliated hairs on the apical part of PES exhibited apparently differences among species. Some problematic species can be differed distinctly by analyzing the combination these elytra with PES ultrastructure:P. denticulatus is not a synonym of P. crenulidens as Benesh wrote in1960, but two valid species due to their remarkably different affiliated hairs; elytra and PES ultrastructure between yellow spot and whole dark elytra individuals of N. delicatus showed very highly similarity, which confirmed that two different splash types are actually existed; O. cuvera sinensis and O. cuvera fallaciosus could be two good species because the shapes and adherent direction between their affiliated hairs are conspicuously different.
引文
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