微红梢斑螟蛀道节肢动物结构及优势种种群分化研究
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摘要
微红梢斑螟（Dioryctria rebulla）是我国重要的历史性松梢害虫。该害虫偏好危害松树幼林，由于发生世代复杂，加之幼虫期隐蔽危害，目前微红梢斑螟的防控仍存在较大的技术瓶颈。本课题以我国微红梢斑螟蛀道伴生生物为研究对象，通过对蛀道内伴生生物分布区系、种类的调查以及诸种类生态位关系的研究，优势种群的筛选以及种类鉴定，发现松梢隐翅虫（Placusa pinearum Gao, Ji, Liu, sp. nov）在时空生态位上与微红梢斑螟具有高度的跟随性和重叠性，可作为载体昆虫携带病原菌防治微红梢斑螟的危害。
在天敌（媒介）昆虫引种和生物防治实践中，首要关注的问题是在不同害虫分布区选择合适的天敌昆虫种源。一般而言，天敌昆虫或生物菌存在明显的地理遗传分化时，综合考虑本地天敌种群对本地气候及寄主等的适应性、定殖扩散能力，一般选择本地的天敌种群以提高防治效率；相反，如果天敌昆虫不同地理种群遗传分化很小，则可在天敌种群基因库内任意选择虫源。基于松梢隐翅虫广泛分布的特点，本研究利用形态标记对不同地理种群的松梢隐翅虫形态分化进行分析，同时，结合分子谱系地理学原理和方法，探讨了松梢隐翅虫种群现有遗传多样性和遗传结构形成的历史过程。目的在于为应用载体昆虫携带病原生物防治微红梢斑螟等钻蛀害虫提供依据，开发林木钻蛀性害虫生物防治的新途径。主要研究内容及结果如下：
1、对南京及周边地区4个不同立地类型的微红梢斑螟危害松林情况进行了调查。林间调查发现微红梢斑螟偏好危害10年生以下的松树幼林。微红梢斑螟对同一松树品种的危害程度与林分类型有较密切关系，林分受害程度依次为：人工纯林>天然更新与人工种植混交林>天然更新林。微红梢斑螟的危害程度与松树树种的关系为：火炬松>马尾松>黑松。
2、首次系统调查了我国不同地区微红梢斑螟蛀道生物区系。微红梢斑螟蛀道节肢动物类群极为丰富。主要类群隶属为3纲9目13科，其中弹尾纲1目1科，为弹尾目跳虫科（Poduridae）；蛛形纲有蜘蛛目和蜱螨目2目2科：分别为螨科（Acarina）和微蛛科（Erigonidae）；昆虫纲6目10科，分别为鞘翅目隐翅虫科（Staphylinidae）、锯谷盗科（Silvanidae）、象甲科（Curculionidae）、小蠹科（Scolytidae）；缨翅目蓟马科（Thripidae）；革翅目蠼螋科（Labiduridae）；膜翅目蚁科（Formicidae）和茧蜂科（Braconidae）；半翅目蝽科（Pentatomidae）；啮虫目啮虫科（Psocidae）。从种群数量及分布特征来看，螨科和隐翅虫科是广布种也是优势种群，另外，从生态位重叠值看，隐翅虫科在时间和空间上与微红梢斑螟存在紧密的共栖关系，提示该甲虫可以作为载体昆虫携带病原菌开展微红梢斑螟的生物防治。
3、发现并报道微红梢斑螟蛀道优势种群前角隐翅虫亚科Placusa属下的一个新种。采集自我国8个不同地区的微红梢斑螟蛀道隐翅虫的44个标本进行观察描述。通过生殖器官解剖、制作玻片、显微拍照，绘制分类特征图，对标本口器细部等特征采用扫描电子显微镜进行观察拍照，发现该隐翅虫在雄性生殖器形态、雌虫受精囊，第8腹节背板和腹板等关键形态特征等与国外已报道种类存在明显的差异，故确定其为一个新种，定名为松梢隐翅虫Placusa pinearum Gao,Ji, Liu, sp. nov。
4、松梢隐翅虫种群的形态分化研究。采集了我国云南、四川、湖南、广东、安徽、江苏和陕西等地区的14个采样点的不同种群，对其头宽、头长、鞘翅宽、鞘翅长、前胸背板宽、前胸背板长、体长以及鞘翅颜色等形态变量进行统计，从鞘翅颜色观察发现云南、四川地区的样品具有黑色的鞘翅，而其他地区的则均为黄色；利用SPSS软件进行聚类分析表明，分布在我国云南四川地区的标本与其他地区的样本存在较为明显的聚类分化，其中云南地区分化最为明显，四川与华东和华南地区亦存在较大分化。
5、利用线粒体mtDNA COI、16S等分子标记对我国14个不同采样点的松梢隐翅虫种群的遗传多样性以及遗传分化进行了研究。结果显示来自西南地区种群单倍型多样性和核苷酸多样性值最低，而华东等地区的种群遗传多样性比较高；实验获得线粒体mtDNACOI、16S基因序列56条和37条，分别产生19条和6条单倍型序列。利用Network以及贝叶斯进化树分析发现，华东地区的种群产生了比较显著的遗传分化，根据遗传分化指数Fst以及共享单倍型的分布特征发现云南四川地区的种群与华东、华南等地区存在显著的遗传分化。
6、形态分化（表型）与遗传分化的关系。从我国不同地理分布地区松梢隐翅虫的形态变异聚类结果看，西南地区与我国华东、华南等地区的种群有明显的形态分化关系，再从其遗传分化的角度看，尽管整体上我国松梢隐翅虫种群并未形成清晰的遗传分化结构，但从共享单倍型的分布、遗传分化指数以及系统进化树等分析结果看，西南地区与华东、华南等地区种群存在着显著的遗传分化，这说明总体上，我国松梢隐翅虫种群在形态分化和遗传分化上具有一致性。即在缺少分子生物学数据的情况下，只要我们获得昆虫种群可靠的形态地理分化关系，就可以大致判断相应的遗传分化特征。
7、初步分析了我国松梢隐翅虫种群遗传分化的原因。将我国14个地区的松梢隐翅虫种群分别按照地理区域和寄主松树的差别进行分组， AMOVA统计结果显示绝大部分的遗传变异产生在组群内的不同种群之间，而仅有少部分的遗传变异产生在组群之间，显示我国不同地区的松梢隐翅虫种群并未形成明显的遗传分化地理结构；Mantel test检验结果显示种群之间的遗传分化与地理距离并未存在显著的相关性。
8、种群历史分析。利用贝叶斯分子钟模型，以昆虫的线粒体mtDNA COI基因碱基突变率估算出在距今大约20000年前，我国松梢隐翅虫种群曾发生过一次明显的种群扩张事件，单倍型网络图结构呈星状结构以及中性检验Fu’s结果所提供的证据也都支持种群发生过种群扩张。来自西南地区种群的单倍型多样性和核酸多样性均显著低于华东地区和华南地区的种群，表明我国云南四川地区的种群可能为新定殖的种群或发生过瓶颈效应，推测西南地区的种群可能是由华东或者华南地区迁移形成的种群，结合云贵采样地位于横断山脉的地理阻隔，最终进而形成目前的遗传分化格局。
Dioryctria rubella is the most notorious pest boring damage to pine shoot and causes greateconomic losses in China. Unfortunately, controlling the pine shoot moth D. rubella is difficultdue to its concealed habitat doubled with overlapping of generations. Our studies on thearthropod community structure inside pine shoot tunnels have revealed that many arthropodspecies live in the tunnels of D. rubella. A species of Staphylinidae (identified as Placusapinearum) has been found most frequently inhabit in the pine shoot tunnel, we also found that ahigh degree of niche overlap (co-occurrence) between the pine shoot moth D. rubella and thebeetle P. pinearum, which triggered our interest as to whether this beetle species can be used asan effective vector to transport biological control agents for controlling pine shoot moths.
To develop biological control programs, selection of suitable source colony from naturalenemy population must be focused firstly, because abundant genetic variation has beenconsidered an important factor for successful colonization and establishment of exotic biologicalcontrol agents. In general speaking, those well-adapted biotypes can expand its geographicranges or match to local climate conditions and colonize successfully in a new habitat. Especially,if the genetic divergence of natural enemy population or biological control agents from differentgeographic region is very weak we can select any source colony in the species gene pool.Conversely, if the genetic divergence is distinct, then, selection of local source colony may beappropriate.
Considering the beetle P. pinearum is widely distributed in China, it is essential tounderstand the intraspecific differentiation, here, we examined P. pinearum collected from14different geographical populations in China, and analyzed the genetic diversity, genetic structureand phylogeographical relationship among the samples by using morphological and molecularmarkers. The aim of this study is to provide a reliable foundation for application this beetle as avector insect and to pave a new approach for biological controlling of those pine shoot borer. Themain results are as follows:
1) The investigations on the rate of damaged pine tree in four sites with different conditionsand forest types around Nanjing areas were carried out. The results showed that D. rubella couldcause severer damage to the artificial pure plantations than the natural regeneration forest andcould cause severer damage to P. taeda than P. massoniana than P. thunbergii. There existsignificant negative linear relationship between diameter at breast height and the rate of damagedshoot of P. massoniana.
2) An investigation was made on the population composition and niche of arthropodcommunity in the pine shoot tunnel bored by D. rubella. The populations of the arthropodcommunity were identified as3classe(sCollembola，Insecta，Arachnida),9order(sCollembola， Aranea, Acarina, Dermaptera, Psocoptera, Hemiptera, Thysanoptera, Coleoptera andHymenoptera)and13families(Poduridae, Acarina, Erigonidae, Staphylinidae, Silvanidae,Curculionidae, Scolytidae, Thripidae, Labiduridae, Formicidae, Braconidae, Pentatomidae,Psocidae). Among the13families, Acarina and Staphylinidae were the dominant populations.The spatiotemporal niche overlap calculated by Levins and Pianka formula was the largestbetween D. rubella and Staphylinidae, indicating their closest relationship. This informationpresented here could shed a light on using the specie in Staphylinidae as an effective vector totransport biological control agents for controlling pine shoot moths.
3) The morphological characteristics of individuals of P. pinearum from14sampling sitescovering Jiangsu、Anhui、Hunan、Guangdong、Sichuan、Shannxi and Yunnan province in Chinawere observed with the help of optical microscope and scanning electron microscope (SEM). Thebeetle P. pinearum was identified as a new specie by the following combination of characters:male and female tergite8with one long median projection and two elongate lateral teeth;flagellum coiled about2times in bulbus; and spermatheca with small and spherical capsule,median portion of spermathecal stem twisted and φ-shaped. The morphological characteristics ofP. pinearum are mentioned above are different in form from the other recorded species. Thespecific name is, Placusa pinearum Gao, Ji, Liu, sp. nov, derived from the unique habitat, wherethe type species was collected.
4) Seven measurements were obtained from87individuals of P. pinearum with an ocularmicrometer under the stereomicroscope at a magnification of40×or100×. These measurementsincluded head width (HW), head length (HL), pronotum width (PW), pronotum length (PL),elytra width (EW), elytra length (EL), and body length (BL). In addition, we found that the elytracolor of the specimens collected from YNKM and SCLS was black, whereas that of specimenscollected from other sampling sites was unanimously bronze yellow. The dendrogramconstructed with the UPGMA methods using Euclidean distances indicated that there was amorphological divergence of the populations in P. pinearum between southwestern and otherthree geographic regions (central, northwestern and eastern region).
5) A portion of the mitochondrial cyochrome oxidase subunit I gene (965-nucleotidesequence) was obtained from56individuals and19haplotypes were identified based on28polymorphic sites in this region, and16S gene sequence was also obtained from37individualsand7haplotypes were identified based on6polymorphic sites in this region. The populationsfrom the southwestern region possessed the lowest nucleotide diversity (0.000622) and haplotypediversity (0.600), whereas those from the southern region possessed the highest nucleotidediversity (0.006390) and those from the northwestern region possessed the highest haplotypediversity (1.000). Our study showed that the significant genetic divergence among the differentpopulations in eastern China, meanwhile, absence of shared haplotypes, coupled with high Fstvalues, demonstrated significant genetic divergence between the populations from southwest andother four main geographical regions (central, southern northwestern and eastern region). Theresults of the genetic character of P. pinearum populations mentioned above can also besupported by the analysis using combining mtDNA16S with COI marker.
6)The relationship between morphological and genetic divergence was compared in this study,morphology-based tree shows that there was a morphological divergence of the populations in P.pinearum between southwestern and other three geographic regions. The median-joining networkanalysis indicated the absence of shared haplotype between the southwestern and the othergeographical regions. In addition, the pairwise Fst value between the southwestern and the otherswas significantly high, indicative of weak gene flow between these regions. It suggests thatmorphological divergence is generally in congruence with genetic divergence in P. pinearumpopulations. These findings also suggest that phenotypic traits could serve as a valid criterion forassessing the degree of genetic divergence in P. pinearum populations, especially whenmolecular data are not available.7) The genetic structure of population subdivision was examined by the analysis of molecularvariance (AMOVA), after the populations were grouped either by geographic locations or by hostspecies. The AMOVA results from our study showed that the majority of molecular variationwas within populations, whereas only a small part was among groups, it was conformed thatthere was no distinct geographical structure of genetic variation among different P. pinearumgroups. Moreover, the mantel test showed that there was no significant correlation between theFst values and geographical distances (km)(r=0.219, P=0.255>0.05).
8) The demographic history of P. pinearum was inferred based on the Bayesian skyline plot,without any assumption of a particular demographic model. In the absence of a suitable intrinsiccalibration, we can apply the approximate2%per million year divergence rate for insect mtDNACOI gene. The Bayesian skyline analysis demonstrates that the P. pinearum populations in Chinahad undergone a major sudden population expansion starting from approximately around20,000years ago, which was also confirmed by the Fu’s test. The low genetic diversity of thesouthwestern populations suggested that it was colonized very recently and might becomeisolated either through dispersal from the eastern or south+central regions or by undergoingbottlenecks due to population size fluctuations. We infer that genetic divergence between thesouthwestern and other region populations might be explained by the consequence of topographicbarriers; that is, the southwestern samples sites (SCLS and YNKM) were in the HengduanMountains, the second largest mountain in China, in contrast to the hills or platforms in theeastern, central, or southern regions.
在天敌（媒介）昆虫引种和生物防治实践中，首要关注的问题是在不同害虫分布区选择合适的天敌昆虫种源。一般而言，天敌昆虫或生物菌存在明显的地理遗传分化时，综合考虑本地天敌种群对本地气候及寄主等的适应性、定殖扩散能力，一般选择本地的天敌种群以提高防治效率；相反，如果天敌昆虫不同地理种群遗传分化很小，则可在天敌种群基因库内任意选择虫源。基于松梢隐翅虫广泛分布的特点，本研究利用形态标记对不同地理种群的松梢隐翅虫形态分化进行分析，同时，结合分子谱系地理学原理和方法，探讨了松梢隐翅虫种群现有遗传多样性和遗传结构形成的历史过程。目的在于为应用载体昆虫携带病原生物防治微红梢斑螟等钻蛀害虫提供依据，开发林木钻蛀性害虫生物防治的新途径。主要研究内容及结果如下：
1、对南京及周边地区4个不同立地类型的微红梢斑螟危害松林情况进行了调查。林间调查发现微红梢斑螟偏好危害10年生以下的松树幼林。微红梢斑螟对同一松树品种的危害程度与林分类型有较密切关系，林分受害程度依次为：人工纯林>天然更新与人工种植混交林>天然更新林。微红梢斑螟的危害程度与松树树种的关系为：火炬松>马尾松>黑松。
2、首次系统调查了我国不同地区微红梢斑螟蛀道生物区系。微红梢斑螟蛀道节肢动物类群极为丰富。主要类群隶属为3纲9目13科，其中弹尾纲1目1科，为弹尾目跳虫科（Poduridae）；蛛形纲有蜘蛛目和蜱螨目2目2科：分别为螨科（Acarina）和微蛛科（Erigonidae）；昆虫纲6目10科，分别为鞘翅目隐翅虫科（Staphylinidae）、锯谷盗科（Silvanidae）、象甲科（Curculionidae）、小蠹科（Scolytidae）；缨翅目蓟马科（Thripidae）；革翅目蠼螋科（Labiduridae）；膜翅目蚁科（Formicidae）和茧蜂科（Braconidae）；半翅目蝽科（Pentatomidae）；啮虫目啮虫科（Psocidae）。从种群数量及分布特征来看，螨科和隐翅虫科是广布种也是优势种群，另外，从生态位重叠值看，隐翅虫科在时间和空间上与微红梢斑螟存在紧密的共栖关系，提示该甲虫可以作为载体昆虫携带病原菌开展微红梢斑螟的生物防治。
3、发现并报道微红梢斑螟蛀道优势种群前角隐翅虫亚科Placusa属下的一个新种。采集自我国8个不同地区的微红梢斑螟蛀道隐翅虫的44个标本进行观察描述。通过生殖器官解剖、制作玻片、显微拍照，绘制分类特征图，对标本口器细部等特征采用扫描电子显微镜进行观察拍照，发现该隐翅虫在雄性生殖器形态、雌虫受精囊，第8腹节背板和腹板等关键形态特征等与国外已报道种类存在明显的差异，故确定其为一个新种，定名为松梢隐翅虫Placusa pinearum Gao,Ji, Liu, sp. nov。
4、松梢隐翅虫种群的形态分化研究。采集了我国云南、四川、湖南、广东、安徽、江苏和陕西等地区的14个采样点的不同种群，对其头宽、头长、鞘翅宽、鞘翅长、前胸背板宽、前胸背板长、体长以及鞘翅颜色等形态变量进行统计，从鞘翅颜色观察发现云南、四川地区的样品具有黑色的鞘翅，而其他地区的则均为黄色；利用SPSS软件进行聚类分析表明，分布在我国云南四川地区的标本与其他地区的样本存在较为明显的聚类分化，其中云南地区分化最为明显，四川与华东和华南地区亦存在较大分化。
5、利用线粒体mtDNA COI、16S等分子标记对我国14个不同采样点的松梢隐翅虫种群的遗传多样性以及遗传分化进行了研究。结果显示来自西南地区种群单倍型多样性和核苷酸多样性值最低，而华东等地区的种群遗传多样性比较高；实验获得线粒体mtDNACOI、16S基因序列56条和37条，分别产生19条和6条单倍型序列。利用Network以及贝叶斯进化树分析发现，华东地区的种群产生了比较显著的遗传分化，根据遗传分化指数Fst以及共享单倍型的分布特征发现云南四川地区的种群与华东、华南等地区存在显著的遗传分化。
6、形态分化（表型）与遗传分化的关系。从我国不同地理分布地区松梢隐翅虫的形态变异聚类结果看，西南地区与我国华东、华南等地区的种群有明显的形态分化关系，再从其遗传分化的角度看，尽管整体上我国松梢隐翅虫种群并未形成清晰的遗传分化结构，但从共享单倍型的分布、遗传分化指数以及系统进化树等分析结果看，西南地区与华东、华南等地区种群存在着显著的遗传分化，这说明总体上，我国松梢隐翅虫种群在形态分化和遗传分化上具有一致性。即在缺少分子生物学数据的情况下，只要我们获得昆虫种群可靠的形态地理分化关系，就可以大致判断相应的遗传分化特征。
7、初步分析了我国松梢隐翅虫种群遗传分化的原因。将我国14个地区的松梢隐翅虫种群分别按照地理区域和寄主松树的差别进行分组， AMOVA统计结果显示绝大部分的遗传变异产生在组群内的不同种群之间，而仅有少部分的遗传变异产生在组群之间，显示我国不同地区的松梢隐翅虫种群并未形成明显的遗传分化地理结构；Mantel test检验结果显示种群之间的遗传分化与地理距离并未存在显著的相关性。
8、种群历史分析。利用贝叶斯分子钟模型，以昆虫的线粒体mtDNA COI基因碱基突变率估算出在距今大约20000年前，我国松梢隐翅虫种群曾发生过一次明显的种群扩张事件，单倍型网络图结构呈星状结构以及中性检验Fu’s结果所提供的证据也都支持种群发生过种群扩张。来自西南地区种群的单倍型多样性和核酸多样性均显著低于华东地区和华南地区的种群，表明我国云南四川地区的种群可能为新定殖的种群或发生过瓶颈效应，推测西南地区的种群可能是由华东或者华南地区迁移形成的种群，结合云贵采样地位于横断山脉的地理阻隔，最终进而形成目前的遗传分化格局。
Dioryctria rubella is the most notorious pest boring damage to pine shoot and causes greateconomic losses in China. Unfortunately, controlling the pine shoot moth D. rubella is difficultdue to its concealed habitat doubled with overlapping of generations. Our studies on thearthropod community structure inside pine shoot tunnels have revealed that many arthropodspecies live in the tunnels of D. rubella. A species of Staphylinidae (identified as Placusapinearum) has been found most frequently inhabit in the pine shoot tunnel, we also found that ahigh degree of niche overlap (co-occurrence) between the pine shoot moth D. rubella and thebeetle P. pinearum, which triggered our interest as to whether this beetle species can be used asan effective vector to transport biological control agents for controlling pine shoot moths.
To develop biological control programs, selection of suitable source colony from naturalenemy population must be focused firstly, because abundant genetic variation has beenconsidered an important factor for successful colonization and establishment of exotic biologicalcontrol agents. In general speaking, those well-adapted biotypes can expand its geographicranges or match to local climate conditions and colonize successfully in a new habitat. Especially,if the genetic divergence of natural enemy population or biological control agents from differentgeographic region is very weak we can select any source colony in the species gene pool.Conversely, if the genetic divergence is distinct, then, selection of local source colony may beappropriate.
Considering the beetle P. pinearum is widely distributed in China, it is essential tounderstand the intraspecific differentiation, here, we examined P. pinearum collected from14different geographical populations in China, and analyzed the genetic diversity, genetic structureand phylogeographical relationship among the samples by using morphological and molecularmarkers. The aim of this study is to provide a reliable foundation for application this beetle as avector insect and to pave a new approach for biological controlling of those pine shoot borer. Themain results are as follows:
1) The investigations on the rate of damaged pine tree in four sites with different conditionsand forest types around Nanjing areas were carried out. The results showed that D. rubella couldcause severer damage to the artificial pure plantations than the natural regeneration forest andcould cause severer damage to P. taeda than P. massoniana than P. thunbergii. There existsignificant negative linear relationship between diameter at breast height and the rate of damagedshoot of P. massoniana.
2) An investigation was made on the population composition and niche of arthropodcommunity in the pine shoot tunnel bored by D. rubella. The populations of the arthropodcommunity were identified as3classe(sCollembola，Insecta，Arachnida),9order(sCollembola， Aranea, Acarina, Dermaptera, Psocoptera, Hemiptera, Thysanoptera, Coleoptera andHymenoptera)and13families(Poduridae, Acarina, Erigonidae, Staphylinidae, Silvanidae,Curculionidae, Scolytidae, Thripidae, Labiduridae, Formicidae, Braconidae, Pentatomidae,Psocidae). Among the13families, Acarina and Staphylinidae were the dominant populations.The spatiotemporal niche overlap calculated by Levins and Pianka formula was the largestbetween D. rubella and Staphylinidae, indicating their closest relationship. This informationpresented here could shed a light on using the specie in Staphylinidae as an effective vector totransport biological control agents for controlling pine shoot moths.
3) The morphological characteristics of individuals of P. pinearum from14sampling sitescovering Jiangsu、Anhui、Hunan、Guangdong、Sichuan、Shannxi and Yunnan province in Chinawere observed with the help of optical microscope and scanning electron microscope (SEM). Thebeetle P. pinearum was identified as a new specie by the following combination of characters:male and female tergite8with one long median projection and two elongate lateral teeth;flagellum coiled about2times in bulbus; and spermatheca with small and spherical capsule,median portion of spermathecal stem twisted and φ-shaped. The morphological characteristics ofP. pinearum are mentioned above are different in form from the other recorded species. Thespecific name is, Placusa pinearum Gao, Ji, Liu, sp. nov, derived from the unique habitat, wherethe type species was collected.
4) Seven measurements were obtained from87individuals of P. pinearum with an ocularmicrometer under the stereomicroscope at a magnification of40×or100×. These measurementsincluded head width (HW), head length (HL), pronotum width (PW), pronotum length (PL),elytra width (EW), elytra length (EL), and body length (BL). In addition, we found that the elytracolor of the specimens collected from YNKM and SCLS was black, whereas that of specimenscollected from other sampling sites was unanimously bronze yellow. The dendrogramconstructed with the UPGMA methods using Euclidean distances indicated that there was amorphological divergence of the populations in P. pinearum between southwestern and otherthree geographic regions (central, northwestern and eastern region).
5) A portion of the mitochondrial cyochrome oxidase subunit I gene (965-nucleotidesequence) was obtained from56individuals and19haplotypes were identified based on28polymorphic sites in this region, and16S gene sequence was also obtained from37individualsand7haplotypes were identified based on6polymorphic sites in this region. The populationsfrom the southwestern region possessed the lowest nucleotide diversity (0.000622) and haplotypediversity (0.600), whereas those from the southern region possessed the highest nucleotidediversity (0.006390) and those from the northwestern region possessed the highest haplotypediversity (1.000). Our study showed that the significant genetic divergence among the differentpopulations in eastern China, meanwhile, absence of shared haplotypes, coupled with high Fstvalues, demonstrated significant genetic divergence between the populations from southwest andother four main geographical regions (central, southern northwestern and eastern region). Theresults of the genetic character of P. pinearum populations mentioned above can also besupported by the analysis using combining mtDNA16S with COI marker.
6)The relationship between morphological and genetic divergence was compared in this study,morphology-based tree shows that there was a morphological divergence of the populations in P.pinearum between southwestern and other three geographic regions. The median-joining networkanalysis indicated the absence of shared haplotype between the southwestern and the othergeographical regions. In addition, the pairwise Fst value between the southwestern and the otherswas significantly high, indicative of weak gene flow between these regions. It suggests thatmorphological divergence is generally in congruence with genetic divergence in P. pinearumpopulations. These findings also suggest that phenotypic traits could serve as a valid criterion forassessing the degree of genetic divergence in P. pinearum populations, especially whenmolecular data are not available.7) The genetic structure of population subdivision was examined by the analysis of molecularvariance (AMOVA), after the populations were grouped either by geographic locations or by hostspecies. The AMOVA results from our study showed that the majority of molecular variationwas within populations, whereas only a small part was among groups, it was conformed thatthere was no distinct geographical structure of genetic variation among different P. pinearumgroups. Moreover, the mantel test showed that there was no significant correlation between theFst values and geographical distances (km)(r=0.219, P=0.255>0.05).
8) The demographic history of P. pinearum was inferred based on the Bayesian skyline plot,without any assumption of a particular demographic model. In the absence of a suitable intrinsiccalibration, we can apply the approximate2%per million year divergence rate for insect mtDNACOI gene. The Bayesian skyline analysis demonstrates that the P. pinearum populations in Chinahad undergone a major sudden population expansion starting from approximately around20,000years ago, which was also confirmed by the Fu’s test. The low genetic diversity of thesouthwestern populations suggested that it was colonized very recently and might becomeisolated either through dispersal from the eastern or south+central regions or by undergoingbottlenecks due to population size fluctuations. We infer that genetic divergence between thesouthwestern and other region populations might be explained by the consequence of topographicbarriers; that is, the southwestern samples sites (SCLS and YNKM) were in the HengduanMountains, the second largest mountain in China, in contrast to the hills or platforms in theeastern, central, or southern regions.
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