中国筒天牛属分类研究（鞘翅目：天牛科：沟胫天牛亚科）
摘要
筒天牛属Oberea Dejean,1835隶属于昆虫纲Insecta,鞘翅目Coleoptera,天牛科Cerambycidae,沟胫天牛亚科Lamiinae。目前,全世界已报道筒天牛属2亚属300余种(亚种)。筒天牛属昆虫分布于除新热带区外的世界各大动物地理区系,部分种类具有重要经济意义。
本文对该属昆虫进行了较为系统的分类研究。文中详细介绍了筒天牛属的经济重要性、研究历史和现状、外部形态特征；在检视了大量的标本,尤其是模式标本的基础上,系统整理、订正和记述了中国分布的筒天牛属昆虫种类；对其进行了比较形态学研究,筛选出具有分类学意义的特征；基于外部形态特征对中国筒天牛属系统发育关系进行了初探；提取了部分种类的分子条形码序列(COI)；并进行了区系分析。研究的主要结果如下：
1.分类研究
本研究查阅了大量文献资料,包括中国分布筒天牛属77种(亚种)的原始描述,检视了700余号标本,其中包括60余种模式标本,模式标本(及照片)检视率达80%以上。在此基础上整理、订正和记述了中国分布的筒天牛属昆虫2亚属,77种(亚种),其中包括1新种,3个中国新记录种,并提供了筒天牛73种成虫外部形态图片和51种雄性外生殖器特征图。其中：
(1)发现1个新种:
拟台湾筒天牛Oberea pseudoformosana Li, Cuccodoro&Chen,2014(已发表)；
(2)发现3个中国新记录种：
褐胫马来筒天牛O. subabdominalis Breuning,1962
褐胫筒天牛O. angustata Pic1923
瞳筒天牛O. pupillata (Gyllenhal,1817)
(3)订正6个新异名：
O. taiwana Matsushita,1933=O. taihokuensis Breuning,1962syn. nov.
O. angustata Pic,1923=O. ohbayashi Kurihara2009syn. nov.
O. clara Pascoe,1866=O. fuscipennis ssp. perakensis Breuning,1962syn. nov.
O. hebescens Bates,1873=O. diversimembris Pic,1923syn. nov.
O. diversipes Pic,1919=O. fuscipennis ssp. fairmairei Breuning,1962syn. nov.
O. walkeri Gahan,1894=O. bicoloritarsis v. subparallela Pic,1928, syn. nov.
(4)提出3个新组合：
Obereopsis subsericea (Breuning,1960) comb. nov.=Oberea subsericea Breuning,1960
Obereopsis binotaticollis chinensis comb. nov.=Oberea griseopennis chinensis Breuning,1982
Obereopsis binotaticollis ichangensis (Breuning,1969) comb. nov.=Oberea griseopennis ichangensis Breuning,1969
(5)恢复了7个种的地位：
黄黑筒天牛O. flavescens Breuning,1947stat. rev.
华东筒天牛O. sylvia Pascoe,1858stat. rev.
和平筒天牛O. binhana Pic,1923stat. rev.
费氏暗翅筒天牛O. diversipes Pic,1919stat. rev.
褐角筒天牛O. toi Gressitt,1939stat. rev.
黄翅筒天牛O. infratestacea Pic,1936stat. rev.
西藏筒天牛O. thibetana Pic,1916stat. rev.
(6)订正了筒天牛属中的79个亚种下名称,具体名称见论文第四章。
2.比较形态学研究
根据文献资料选取了筒天牛属昆虫40个形态特征进行比较研究,筛选出了27个具有分类学意义的特征：
头部具有分类学意义的特征：头部颜色、额长宽比、复眼下叶与颊长之比、触角与体长之比、触角第3、4节之比、触角柄节是否具有纵沟等6个特征。
胸部和腹部具有分类学意义的特征：前胸长宽比、鞘翅末端形状、缘角是否具刺、鞘翅刻点的大小及是否排列整齐、后足腿节的长度、后足胫节与跗节之比、鞘翅中缝是否被丝状绒毛、第1、2可见腹节是否被丝状绒毛、第1可见腹节是否具有指状突起等11个特征。
雄性外生殖器具有分类意义的特征有：第8腹节背板的形态、阳基侧突形态、内囊端部骨针的数量和形态、长骨针与短骨针长度之比、长骨针和短骨针是否对称,长骨针端部是否相连,短骨针是否基部至少相连等10个特征。
其中,额长宽比、后足胫节与跗节之比、雄性第8腹节背板的形态、阳基侧突的形态4个特征,首次用于该属的分类鉴定。根据比较形态学研究结果,完善了分类学部分的研究结果,对部分种类进行了重新描述和修订,编制了筒天牛属分亚属、亚属分种的检索表。
3.系统发育研究
本研究选取了小筒天牛族的3属3种天牛作为外群,利用PAUP4.0,基于比较形态学筛选的26个成虫形态特征,使用邻接法对中国55种筒天牛属昆虫的系统发育关系进行了支序分析,得到的NJ树显示：
(1)筒天牛属昆虫的单系性受到质疑,外群瘤筒天牛属Linda与内群聚到一起,说明筒天牛属Oberea是并系群；
(2)筒天牛亚属Oberea和暗口筒天牛亚属Amaurostoma在支序图中并未分别聚成一支,表明Amaurostoma应作为筒天牛属的次异名而不是亚属更为合理；
(3)支序图的分支与雄性外生殖器内囊骨化物的形态具有密切联系：滇筒天牛O. yunnanensis Breuning作为单独一支首先分出,该种的雄性外生殖器内囊端部仅具1对细长骨针；筒天牛属的其余类群分为两个大支(支系Ⅰ和支系Ⅱ),支系Ⅰ中的25种简天牛的内囊骨化物短骨针基部均分离,支系Ⅱ中的29种内囊骨化物短骨针至少基部相连。因此认为应根据雄性外生殖器的结构对筒天牛属划分亚属或者种团。由于本研究仅涉及中国分布的种类,加之节点支持率较低,目前无法给出更合适的建议。有待今后对世界筒天牛进行研究时增加种类、结合形态和分子数据重建筒天牛属的分类系统。
4.DNA条形码研究
利用通用引物LCO1490/HCO2198扩增筒天牛属6种18号标本的COI序列,得到长度为658bp的片段,结合GenBank数据库下载的5种筒天牛COI序列,以K2P模型计算种内及种间遗传距离,并利用NJ法和ML法构建系统发育树,结果显示：
(1)种内遗传距离为0-2.04%,种间遗传距离为5.97%-12.74%,种间遗传距离明显大于种内遗传距离；基于DNA条形码序列,利用邻接法和最大似然法,构建了系统发育树,同一个种聚成一小支,且节点支持率均为100%,遗传距离和系统发育树的结果都说明该片段可以作为分析和鉴别筒天牛属物种依据。
(2)利用分子条形码数据验证了分类学部分的研究结果。在分类学部分,本研究根据雄性外生殖器等特征恢复了暗翅筒天牛O.fuscipennis (Chevrolat)的两个近似种费氏暗翅筒天牛O. diversipes Pic和黄翅筒天牛O. infratestacea Pic的地位,从DNA条形码数据计算得到的遗传距离和构建的系统发育树结果显示这三个种种间遗传距离明显大于种内遗传距离,且每个种各自聚为一小支,证明这三个种均为独立的有效种；由于这三个种雌虫外部形态和雌性生殖器特征相似,雌虫鉴定较为困难,通过DNA条形码技术可以准确鉴定雌虫。同时,验证了凹尾筒天牛O.walkeri第2可见腹节两侧的黑斑属于种内变异,该特征不能作为鉴定特征。
5.区系分析
根据文献资料和标本采集信息,本研究对我国筒天牛属昆虫种类在世界和中国动物地理区划的分布情况进行了统计,并对其区系分布进行了简要的分析,结果表明：
(1)在世界动物地理区划中,我国筒天牛属昆虫仅分布在东洋界和古北界,且以东洋界占优势。只分布在古北界的仅有11种,占已知种类的14.29%；仅分布在东洋界的共46种,占59.74%；古北界-东洋界分布20种,占25.97%。从亚属水平上分析,暗口筒天牛亚属Amaurostoma仅分布在古北界；而筒天牛亚属Oberea在古北界和东洋界均有分布。
(2)在我国动物地理区划中,华中区的筒天牛属昆虫种类最丰富,共40种,占已知种类的51.95%,其次为华南区,共37种,占48.05%；以后依次为西南区=华中区>青藏区>东北区>蒙新区。筒天牛属昆虫共有25种分布型,其中暗口筒天牛亚属Amaurostoma具有3种分布型：华北区、蒙新区、华北区-蒙新区,每分布型各有1种,分别占总数的1.33%；筒天牛亚属Oberea具有22种分布型,单区分布的种类最多,共41种,占56.94%,单区分布的种类又以华南区和华中区最多,分别为15种和14种,占该亚属的20.83%和19.44%；跨区分布的种类较少,无全国广布种。
(3)中国特有成分丰富,在我国记录分布的77种中,有31个中国特有种,占我国筒天牛属昆虫的40.3%。华南区分布的特有种最多,其次是华中区,蒙新区无特有种。
Genus Oberea Dejean,1835belonging to subfamily Lamminae, family Cerambycidae, order Coleoptera consists of2subgenera, more than300species. Oberea is widely distributed in most zoogeographical realms. Generally, some Oberea species are economically important to agriculture and forestry. The present dissertation introduces the brief research history, general morphology of Oberea. All the species from China are described in details. The taxonomic characters are compared. The phylogenetic study on Oberea is made based on the morphological characters. The sequences of DNA barcode based on COI are extracted. The zoogeography of Oberea is analyzed. The principal results of the present dissertation are listed as follows:
1. Taxonomy
In the taxonomic section,77species (subspecies) of Oberea are recorded and described in details base on more than700specimens, including more than60type specimens. A new species, six new country records, six status resurrected, six new synonym and three new combinations are recorded. The adult habitus of73species and male genitalia of51species are provided.
One new species:
Oberea pseudoformosana Li, Cuccodoro&Chen,2014(published)
Three new recorded species to China:
O. subabdominalis Breuning,1962
O. angustata Pic1923
O.pupillata (Gyllenhal,1817).
Six new synonyms:
O. taiwana Matsushita,1933=O. taihokuensis Breuning,1962syn.nov.
O. angustata Pic,1923=O. ohbayashi Kurihara2009. syn. nov
O. clara Pascoe,1866=O.fuscipennis ssp. perakensis Breuning,1962syn. nov
O. hebescens Bates,1873=O. diversimembris Pic,1923syn. nov
O. diversipes Pic,1919=O. fuscipennis ssp. fairmairei Breuning,1962syn. nov
O. walkeri Gahan,1894=0. bicoloritarsis v. subparallela Pic,1928, syn. nov.
Three new combinations:
Obereopsis subsericea (Breuning,1960) comb. nov.=Oberea subsericea Breuning,1960
Obereopsis binotaticollis chinensis comb. nov.=Oberea griseopennis chinensis Breuning,1982
Obereopsis binotaticollis ichangensis (Breuning,1969) comb. nov.=Oberea griseopennis ichangensis Breuning,1969
Seven status resurrected:
O. flavescens Breuning,1947stat. rev.,
O. sylvia Pascoe,1858stat. rev.
O. binhana Pic,1923stat. rev.
O. diversipes Pic,1919stat. rev.
O. toi Gressitt,1939stat. rev.
O. infratestacea Pic,1936stat.rev.
O. thibetana Pic,1916stat. rev..
Seventy-nine infrasubspecific names are revised.
2. Comparative morphology
The present comparative morphology research is based on40morphological characters of which27taxonomic characters can be used.
There are6taxonomic characters of head:color of head, shape of frons, ratio of lower lobe of eyes and genea, antennal length, third/fourth antennal segment, groove on the dorsal surface of scape,.
There are11taxonomic characters of thorax and abdomen:ratio between length and width of prothorax, elytral apex rounded or truncated, outer elytral angles with or without spine, punctures on the elytral disc, silky pubescence on the elytra, length of hind femora, ratio of hind tibae and tarsis, silky pubescence on the abdominal segments, the finger-like protuding of the first visible abdominal segment of males and color of last visible abdominal segment.
There are10taxonomic characters of male genitalia:shape of tergite VIII, shape of lateral lobes, number, symmetry of sclerity in apical endophallus, ratio between long pairs and short pairs, and so on.
Based on the results of comparative morphology, some species have been redescribed and revised, and the keys to subgenera and species are established.
3. Phylogenetic analysis
The phylogenetic relationship among the55species of Oberea is re-constructed based on the26morphological characters.
The NJ tree shows that Oberea is a paraphyly because outgroup Linda are gathered within Oberea. The relationship of Linda and Oberea need further study.
Species of subgenus Oberea and Amaurostoma are not gathered as independent branch. Thus Amaurostoma should be considered as synonym of Oberea rather than subgenus.
According to the relationship between the branch and characters of male genitalia, the classification system of genus Oberea should be reconstructed based on the characters of male genitalia, especially the rods of endopallus. However, the current research involves in the Chinese species only accounting for only1/6of worldwide species. More species, morphological and molecular data are needed for classification system reconstruciton of Oberea.
4. DNA barcode
The658bp sequence of COI extracted from6species,18specimens of Oberea with the COI sequence of5species from GenBank are analyzed. Model Kimura2-parameter (K2P) is used to analyze the genetic distance. And the results shows that the genetic distance among the same species varies from0-2.04%, while that among different species ranged from5.97%-12.74%. It shows that the inter-specific genetic distance is significantly larger than intra-specific genetic distance. Neighbor-joining phylogenetic tree and ML tree are established based on the COI gene sequences. Both NJ tree and ML tree indicate that the same species was gathered in the same branch, with the bootstrap value of100%. The results suggest that DNA barcoding based on mitochondrial COI gene sequences is applicable in the classification and identification of Oberea species.
In the taxonomic section, the status of O. diversipes Pic and O. infratestacea Pic which are closely related to O. fuscipennis was resurrected based on the morphology of male genitalia. The inter-specific genetic distance is larger than intra-specific genetic distance. And the same species are gather within the same branch. Therefore, O. diversipes Pic and O. infratestacea Pic are good species. The DNA barcode sequence of three specimens of O. walkeri Gahan, two of which have black spots on the second visible abdominal segment are analyzed. The results indicate that three specimen are the same species and the black spots on the second visible abdominal segment is variation of O. walkeri, which cannot be used for identification of Oberea.
5. Zoogeographical distribution
According to the general data of the geographical distribution and collection records, the distribution patterns of Oberea are summarized both world and China scale.
It shows that Chinese Oberea are distributed in Oriental realm and Palearctic realm. And the Oriental realm holds the richest species diversity of Oberea. There are11species are distributed in Palearctic realm, accounting for14.29%. There are46species are distributed in Oriental realm, having the highest proportion of59.74%. There are20species are distributed both in Oriental and Palearctic realm. Species of subgenera Amaurostoma are only distributed in Palearctic realm.
In the Chinese zoogeographic region, Center China Region (40species, accounting for51.95%) holds richest species diversity of Oberea, South China Region comes next and Inner Mongolia-Xinjiang Region holds fewest. There are3region types of subgenera Amaurostoma, while22of subgenera Oberea. There are41species distributed in only one region, accounting for56.94%, of which are mainly distributed in Center China region and South China region. The species distributed over two regions are much less.
There are41species only distributed in one region, accounting for56.94%. And the species distributed more than2regions are much more less than one region distributed species. There are31endemic species from China, most of which are distributed in South China region and Center China region.
本文对该属昆虫进行了较为系统的分类研究。文中详细介绍了筒天牛属的经济重要性、研究历史和现状、外部形态特征；在检视了大量的标本,尤其是模式标本的基础上,系统整理、订正和记述了中国分布的筒天牛属昆虫种类；对其进行了比较形态学研究,筛选出具有分类学意义的特征；基于外部形态特征对中国筒天牛属系统发育关系进行了初探；提取了部分种类的分子条形码序列(COI)；并进行了区系分析。研究的主要结果如下：
1.分类研究
本研究查阅了大量文献资料,包括中国分布筒天牛属77种(亚种)的原始描述,检视了700余号标本,其中包括60余种模式标本,模式标本(及照片)检视率达80%以上。在此基础上整理、订正和记述了中国分布的筒天牛属昆虫2亚属,77种(亚种),其中包括1新种,3个中国新记录种,并提供了筒天牛73种成虫外部形态图片和51种雄性外生殖器特征图。其中：
(1)发现1个新种:
拟台湾筒天牛Oberea pseudoformosana Li, Cuccodoro&Chen,2014(已发表)；
(2)发现3个中国新记录种：
褐胫马来筒天牛O. subabdominalis Breuning,1962
褐胫筒天牛O. angustata Pic1923
瞳筒天牛O. pupillata (Gyllenhal,1817)
(3)订正6个新异名：
O. taiwana Matsushita,1933=O. taihokuensis Breuning,1962syn. nov.
O. angustata Pic,1923=O. ohbayashi Kurihara2009syn. nov.
O. clara Pascoe,1866=O. fuscipennis ssp. perakensis Breuning,1962syn. nov.
O. hebescens Bates,1873=O. diversimembris Pic,1923syn. nov.
O. diversipes Pic,1919=O. fuscipennis ssp. fairmairei Breuning,1962syn. nov.
O. walkeri Gahan,1894=O. bicoloritarsis v. subparallela Pic,1928, syn. nov.
(4)提出3个新组合：
Obereopsis subsericea (Breuning,1960) comb. nov.=Oberea subsericea Breuning,1960
Obereopsis binotaticollis chinensis comb. nov.=Oberea griseopennis chinensis Breuning,1982
Obereopsis binotaticollis ichangensis (Breuning,1969) comb. nov.=Oberea griseopennis ichangensis Breuning,1969
(5)恢复了7个种的地位：
黄黑筒天牛O. flavescens Breuning,1947stat. rev.
华东筒天牛O. sylvia Pascoe,1858stat. rev.
和平筒天牛O. binhana Pic,1923stat. rev.
费氏暗翅筒天牛O. diversipes Pic,1919stat. rev.
褐角筒天牛O. toi Gressitt,1939stat. rev.
黄翅筒天牛O. infratestacea Pic,1936stat. rev.
西藏筒天牛O. thibetana Pic,1916stat. rev.
(6)订正了筒天牛属中的79个亚种下名称,具体名称见论文第四章。
2.比较形态学研究
根据文献资料选取了筒天牛属昆虫40个形态特征进行比较研究,筛选出了27个具有分类学意义的特征：
头部具有分类学意义的特征：头部颜色、额长宽比、复眼下叶与颊长之比、触角与体长之比、触角第3、4节之比、触角柄节是否具有纵沟等6个特征。
胸部和腹部具有分类学意义的特征：前胸长宽比、鞘翅末端形状、缘角是否具刺、鞘翅刻点的大小及是否排列整齐、后足腿节的长度、后足胫节与跗节之比、鞘翅中缝是否被丝状绒毛、第1、2可见腹节是否被丝状绒毛、第1可见腹节是否具有指状突起等11个特征。
雄性外生殖器具有分类意义的特征有：第8腹节背板的形态、阳基侧突形态、内囊端部骨针的数量和形态、长骨针与短骨针长度之比、长骨针和短骨针是否对称,长骨针端部是否相连,短骨针是否基部至少相连等10个特征。
其中,额长宽比、后足胫节与跗节之比、雄性第8腹节背板的形态、阳基侧突的形态4个特征,首次用于该属的分类鉴定。根据比较形态学研究结果,完善了分类学部分的研究结果,对部分种类进行了重新描述和修订,编制了筒天牛属分亚属、亚属分种的检索表。
3.系统发育研究
本研究选取了小筒天牛族的3属3种天牛作为外群,利用PAUP4.0,基于比较形态学筛选的26个成虫形态特征,使用邻接法对中国55种筒天牛属昆虫的系统发育关系进行了支序分析,得到的NJ树显示：
(1)筒天牛属昆虫的单系性受到质疑,外群瘤筒天牛属Linda与内群聚到一起,说明筒天牛属Oberea是并系群；
(2)筒天牛亚属Oberea和暗口筒天牛亚属Amaurostoma在支序图中并未分别聚成一支,表明Amaurostoma应作为筒天牛属的次异名而不是亚属更为合理；
(3)支序图的分支与雄性外生殖器内囊骨化物的形态具有密切联系：滇筒天牛O. yunnanensis Breuning作为单独一支首先分出,该种的雄性外生殖器内囊端部仅具1对细长骨针；筒天牛属的其余类群分为两个大支(支系Ⅰ和支系Ⅱ),支系Ⅰ中的25种简天牛的内囊骨化物短骨针基部均分离,支系Ⅱ中的29种内囊骨化物短骨针至少基部相连。因此认为应根据雄性外生殖器的结构对筒天牛属划分亚属或者种团。由于本研究仅涉及中国分布的种类,加之节点支持率较低,目前无法给出更合适的建议。有待今后对世界筒天牛进行研究时增加种类、结合形态和分子数据重建筒天牛属的分类系统。
4.DNA条形码研究
利用通用引物LCO1490/HCO2198扩增筒天牛属6种18号标本的COI序列,得到长度为658bp的片段,结合GenBank数据库下载的5种筒天牛COI序列,以K2P模型计算种内及种间遗传距离,并利用NJ法和ML法构建系统发育树,结果显示：
(1)种内遗传距离为0-2.04%,种间遗传距离为5.97%-12.74%,种间遗传距离明显大于种内遗传距离；基于DNA条形码序列,利用邻接法和最大似然法,构建了系统发育树,同一个种聚成一小支,且节点支持率均为100%,遗传距离和系统发育树的结果都说明该片段可以作为分析和鉴别筒天牛属物种依据。
(2)利用分子条形码数据验证了分类学部分的研究结果。在分类学部分,本研究根据雄性外生殖器等特征恢复了暗翅筒天牛O.fuscipennis (Chevrolat)的两个近似种费氏暗翅筒天牛O. diversipes Pic和黄翅筒天牛O. infratestacea Pic的地位,从DNA条形码数据计算得到的遗传距离和构建的系统发育树结果显示这三个种种间遗传距离明显大于种内遗传距离,且每个种各自聚为一小支,证明这三个种均为独立的有效种；由于这三个种雌虫外部形态和雌性生殖器特征相似,雌虫鉴定较为困难,通过DNA条形码技术可以准确鉴定雌虫。同时,验证了凹尾筒天牛O.walkeri第2可见腹节两侧的黑斑属于种内变异,该特征不能作为鉴定特征。
5.区系分析
根据文献资料和标本采集信息,本研究对我国筒天牛属昆虫种类在世界和中国动物地理区划的分布情况进行了统计,并对其区系分布进行了简要的分析,结果表明：
(1)在世界动物地理区划中,我国筒天牛属昆虫仅分布在东洋界和古北界,且以东洋界占优势。只分布在古北界的仅有11种,占已知种类的14.29%；仅分布在东洋界的共46种,占59.74%；古北界-东洋界分布20种,占25.97%。从亚属水平上分析,暗口筒天牛亚属Amaurostoma仅分布在古北界；而筒天牛亚属Oberea在古北界和东洋界均有分布。
(2)在我国动物地理区划中,华中区的筒天牛属昆虫种类最丰富,共40种,占已知种类的51.95%,其次为华南区,共37种,占48.05%；以后依次为西南区=华中区>青藏区>东北区>蒙新区。筒天牛属昆虫共有25种分布型,其中暗口筒天牛亚属Amaurostoma具有3种分布型：华北区、蒙新区、华北区-蒙新区,每分布型各有1种,分别占总数的1.33%；筒天牛亚属Oberea具有22种分布型,单区分布的种类最多,共41种,占56.94%,单区分布的种类又以华南区和华中区最多,分别为15种和14种,占该亚属的20.83%和19.44%；跨区分布的种类较少,无全国广布种。
(3)中国特有成分丰富,在我国记录分布的77种中,有31个中国特有种,占我国筒天牛属昆虫的40.3%。华南区分布的特有种最多,其次是华中区,蒙新区无特有种。
Genus Oberea Dejean,1835belonging to subfamily Lamminae, family Cerambycidae, order Coleoptera consists of2subgenera, more than300species. Oberea is widely distributed in most zoogeographical realms. Generally, some Oberea species are economically important to agriculture and forestry. The present dissertation introduces the brief research history, general morphology of Oberea. All the species from China are described in details. The taxonomic characters are compared. The phylogenetic study on Oberea is made based on the morphological characters. The sequences of DNA barcode based on COI are extracted. The zoogeography of Oberea is analyzed. The principal results of the present dissertation are listed as follows:
1. Taxonomy
In the taxonomic section,77species (subspecies) of Oberea are recorded and described in details base on more than700specimens, including more than60type specimens. A new species, six new country records, six status resurrected, six new synonym and three new combinations are recorded. The adult habitus of73species and male genitalia of51species are provided.
One new species:
Oberea pseudoformosana Li, Cuccodoro&Chen,2014(published)
Three new recorded species to China:
O. subabdominalis Breuning,1962
O. angustata Pic1923
O.pupillata (Gyllenhal,1817).
Six new synonyms:
O. taiwana Matsushita,1933=O. taihokuensis Breuning,1962syn.nov.
O. angustata Pic,1923=O. ohbayashi Kurihara2009. syn. nov
O. clara Pascoe,1866=O.fuscipennis ssp. perakensis Breuning,1962syn. nov
O. hebescens Bates,1873=O. diversimembris Pic,1923syn. nov
O. diversipes Pic,1919=O. fuscipennis ssp. fairmairei Breuning,1962syn. nov
O. walkeri Gahan,1894=0. bicoloritarsis v. subparallela Pic,1928, syn. nov.
Three new combinations:
Obereopsis subsericea (Breuning,1960) comb. nov.=Oberea subsericea Breuning,1960
Obereopsis binotaticollis chinensis comb. nov.=Oberea griseopennis chinensis Breuning,1982
Obereopsis binotaticollis ichangensis (Breuning,1969) comb. nov.=Oberea griseopennis ichangensis Breuning,1969
Seven status resurrected:
O. flavescens Breuning,1947stat. rev.,
O. sylvia Pascoe,1858stat. rev.
O. binhana Pic,1923stat. rev.
O. diversipes Pic,1919stat. rev.
O. toi Gressitt,1939stat. rev.
O. infratestacea Pic,1936stat.rev.
O. thibetana Pic,1916stat. rev..
Seventy-nine infrasubspecific names are revised.
2. Comparative morphology
The present comparative morphology research is based on40morphological characters of which27taxonomic characters can be used.
There are6taxonomic characters of head:color of head, shape of frons, ratio of lower lobe of eyes and genea, antennal length, third/fourth antennal segment, groove on the dorsal surface of scape,.
There are11taxonomic characters of thorax and abdomen:ratio between length and width of prothorax, elytral apex rounded or truncated, outer elytral angles with or without spine, punctures on the elytral disc, silky pubescence on the elytra, length of hind femora, ratio of hind tibae and tarsis, silky pubescence on the abdominal segments, the finger-like protuding of the first visible abdominal segment of males and color of last visible abdominal segment.
There are10taxonomic characters of male genitalia:shape of tergite VIII, shape of lateral lobes, number, symmetry of sclerity in apical endophallus, ratio between long pairs and short pairs, and so on.
Based on the results of comparative morphology, some species have been redescribed and revised, and the keys to subgenera and species are established.
3. Phylogenetic analysis
The phylogenetic relationship among the55species of Oberea is re-constructed based on the26morphological characters.
The NJ tree shows that Oberea is a paraphyly because outgroup Linda are gathered within Oberea. The relationship of Linda and Oberea need further study.
Species of subgenus Oberea and Amaurostoma are not gathered as independent branch. Thus Amaurostoma should be considered as synonym of Oberea rather than subgenus.
According to the relationship between the branch and characters of male genitalia, the classification system of genus Oberea should be reconstructed based on the characters of male genitalia, especially the rods of endopallus. However, the current research involves in the Chinese species only accounting for only1/6of worldwide species. More species, morphological and molecular data are needed for classification system reconstruciton of Oberea.
4. DNA barcode
The658bp sequence of COI extracted from6species,18specimens of Oberea with the COI sequence of5species from GenBank are analyzed. Model Kimura2-parameter (K2P) is used to analyze the genetic distance. And the results shows that the genetic distance among the same species varies from0-2.04%, while that among different species ranged from5.97%-12.74%. It shows that the inter-specific genetic distance is significantly larger than intra-specific genetic distance. Neighbor-joining phylogenetic tree and ML tree are established based on the COI gene sequences. Both NJ tree and ML tree indicate that the same species was gathered in the same branch, with the bootstrap value of100%. The results suggest that DNA barcoding based on mitochondrial COI gene sequences is applicable in the classification and identification of Oberea species.
In the taxonomic section, the status of O. diversipes Pic and O. infratestacea Pic which are closely related to O. fuscipennis was resurrected based on the morphology of male genitalia. The inter-specific genetic distance is larger than intra-specific genetic distance. And the same species are gather within the same branch. Therefore, O. diversipes Pic and O. infratestacea Pic are good species. The DNA barcode sequence of three specimens of O. walkeri Gahan, two of which have black spots on the second visible abdominal segment are analyzed. The results indicate that three specimen are the same species and the black spots on the second visible abdominal segment is variation of O. walkeri, which cannot be used for identification of Oberea.
5. Zoogeographical distribution
According to the general data of the geographical distribution and collection records, the distribution patterns of Oberea are summarized both world and China scale.
It shows that Chinese Oberea are distributed in Oriental realm and Palearctic realm. And the Oriental realm holds the richest species diversity of Oberea. There are11species are distributed in Palearctic realm, accounting for14.29%. There are46species are distributed in Oriental realm, having the highest proportion of59.74%. There are20species are distributed both in Oriental and Palearctic realm. Species of subgenera Amaurostoma are only distributed in Palearctic realm.
In the Chinese zoogeographic region, Center China Region (40species, accounting for51.95%) holds richest species diversity of Oberea, South China Region comes next and Inner Mongolia-Xinjiang Region holds fewest. There are3region types of subgenera Amaurostoma, while22of subgenera Oberea. There are41species distributed in only one region, accounting for56.94%, of which are mainly distributed in Center China region and South China region. The species distributed over two regions are much less.
There are41species only distributed in one region, accounting for56.94%. And the species distributed more than2regions are much more less than one region distributed species. There are31endemic species from China, most of which are distributed in South China region and Center China region.
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