中巴法医昆虫种类研究—侧重丽蝇及其分子鉴定
摘要
近些年来越来越多的昆虫学证据被应用于司法实践中,特别是在刑事案件侦破和法庭审判中起着不可低估的作用。全面了解特定地区尸体上昆虫的种类,掌握快速准确的鉴定技术以及环境因素对昆虫种类的影响是正确利用这些昆虫进行法医鉴定的基础,是法医昆虫学研究的重要内容。
本论文分别调查分析了中国和巴基斯坦两国代表性区域法医昆虫的种类与种群演替规律,以及巴基斯坦传统节日—宰牲节对法医昆虫种类等的影响,并对杭州华家池重要类群丽蝇的分子鉴定技术进行了研究。
中国杭州华家池周围的蝇类(双翅目:Diptera)调查结果表明:(1)华家池共有蝇类法医昆虫12种,其中丽蝇科Calliphoridae7种,分别为大头金蝇Chrysomyamegacephala(Fabricius)、肥躯金蝇Ch.pinguis(Walker)、巨尾阿丽蝇Aldrichinagrahami(Aldrich)、丝光绿蝇Lucilia sericata(Meigen)、紫绿蝇L.porphyrina(Walker)、亮绿蝇L.illustris(Meigen)和瘦叶带绿蝇Hemipyrellia,igurriens(Wiedemann);麻蝇科Sarcophagidae一种,即肥须亚麻蝇Parasarcophagacrassipalpis(Macquart);蝇科Muscidae 4种,即家蝇Musca domestica(Linnaeus)、Musca sp.、Fannia sp.、Ophyra sp.;(2)大头金蝇是唯一全年都可以诱集到的种群,而且该种虫口数量占绝对的优势,而紫绿蝇虫口数量最小;(3)2007年1月至2008年12月的调查结果显示,丽蝇科7个种类、肥须亚麻蝇和家蝇的虫口数量在冬季均明显偏低,并且一月份只诱集到了大头金蝇和家蝇两种,二月份诱集到大头金蝇、肥躯金蝇、巨尾阿丽蝇、家蝇等四种,三月份诱集到了大头金蝇、肥躯金蝇、巨尾阿丽蝇、丝光绿蝇、亮绿蝇、肥须亚麻蝇、家蝇等7种;同时,统计分析结果表明虫口数量在这些月份之间没有显著差异,但冬夏种群虫口数量差异显著。
鉴于丽蝇幼期种类在形态上难以区分,本研究以华家池采集的丽蝇为材料,选用来自线粒体基因组的COⅠ、CYTB和ND5以及核糖体基因组的ITS1和ITS2共5个基因片段做分子标记,对丽蝇种类的分子鉴定进行了探索性研究。从7种丽蝇17个标本中扩增得到了75条基因序列并提交到GenBank。结果表明COⅠ、Cytb、ND5和ITS2可作为丽蝇种间分子鉴定的分予标记。ITS1序列可以作为丽蝇同属分子鉴定的分子标记。同时,研究比较了28对蝇(包括丽蝇、家蝇)5个基因位点的种间差异。结果表明:在丽蝇中,(1)ITS1的种内序列差异度为0.29-1.16%,而在种间水平上差异显著,为7.8-57.3%,表明ITS1能够用来进行同属内近缘种类鉴定;(2)ITS2的种间差异度为1.2-19%,表明该序列能够用于种的鉴别,但在姊妹中则可能出错;(3)COI序列的种内差异度在2.2-10.4%之间,也显示在姊妹种的鉴定上可能会错;(4)另一方面,CYTB和ND5序列种间分歧度分别为5.2-15.4%,4.5-12.4%,表明这两个序列均可成功用于种的鉴定。在丽蝇和家蝇组合中,(1)ITS2有30-35.2%的科间序列差异度,从而证明可作为最重要的基因标记用于科级水平的鉴定;(2)COⅠ的序列差异度10.3-13.5%表明在丽蝇科、家蝇科和某些丽蝇科内种类之间(10.4%)不能明显地区分;(3)CYTB序列的科间差异度12.5-16.7%也说明该序列在科技水平上不能明显区分;(4)但ND5序列的科间差异度16-17%则说明在可用于科级水平的鉴定;(5)结果还证明CO1、CYTB、ND5和ITS2等基因在鉴别丽蝇的系统发育关系中也很有用;(6)基于自己获得的基因序列以及GenBank中获得的分子数据,还推算了丽蝇科种类的演化关系。总之,ITS1序列具有低的种内和高的种间差异度,可望用于近缘种的区分;ND5可用于丽蝇不同地理种群的鉴别;ITS2和ND5均表现出科级水平的差异显著,COⅠ和ITS2序列分析表明,丽蝇科和家蝇科大约于20-50 MYA形成不同的进化分支。
在巴基斯坦穆斯林传统节日—宰牲节期间,大量动物尸体残留物抛弃在外,从而引起尸食类昆虫的大量繁殖,而目前尚无脊椎动物在户外垃圾场的降解以及与疾病传播方面的研究报道。本文以巴基斯坦一个典型乡镇为代表,2008年首次对宰牲节期间产生的血坑、肉、皮毛和骨头等14处场所进行了调查,共发现35种昆虫,其中腐肉上面发现的种类和数量最多;首次分析研究了分别来自巴基斯坦D.I.Khan城和Rawalpindi城宰牲节期间红颜金蝇Ch.rufifacies种群的COⅠ、COⅡ和ITS2基因序列,并加以对比分析,发现两个城市不同个体的红颜金蝇的分子序列分析完全相同。同时,在宰牲节期间的调查也发现牲畜交易过程提高了蝇蛆病的传播效率,两个城市的红颜金蝇不同个体分子序列分析没有差异的结果暗示了大量的动物贸易可能会携带并传播一些重要的蝇类,从而导致蝇类种群扩散。另外,还讨论了宰牲节对法医鉴定产生的深远影响。
The present study was aimed at achieving three main goals,to define the urban profile of blowflies,to devise a multiple gene approach for their molecular identification and to examine the association of blowflies and other carrion breeding insects with the Muslim "Festival of sacrifice i.e.Eid-ul-Azha".
Community structure of blowflies was determined in the urban settings of Huajiachi, Hangzhou,eastern China.Standard liver-sodium sulfide baits were placed along various urban facilities,including open sewerage drain,garbage dump sites,mulberry plantation, college building,pig farm and meat-vegetable markets.(1) We collected 12 species in all with nine species of major forensic importance including Chrysomya megacephala,Ch. pinguis,Aldrichina grahami,Lucilia sericata,L.illustris,L.porphyrina,Hemipyrellia ligurriens(Diptera:Calliphoridae),Musca domestica(Muscidae) and Parasarcophaga crassipalpis(Sarcophagidae).(2) ANOVA test statistics revealed significant mean population variation between species(F= 251.670) and months(F= 164.231).(3) Ch. megacephala was the only species observed throughout the study year.This species was the most dominant and abundant group with a population mean of 33±3.919 while L. porphyrina with a mean of 0.27±0.122 was the least represented species.(4) A special assemblage of seven species excluding the blowfly Ch.megacephala and housefly M. domestica were found absent in the January of two consecutive years,i.e.2007 and 2008. In February five species L.sericata,L.porphyrina,L.illustris,H.ligurriens and the flesh fly P.crassipalpis were absent in 2007-08 while in March only L.porphyrina and H. ligurriens were absent in 2007-08.Therefore,winter population means of all nine species were considerably low;January=3.3±0.246,February=1.14±0.472,December=0.25±0.76 and LSD analysis showed no significant difference for these months.However, statistical variation was significant when compared for winter and summer months.
The second phase of our study dealt with the molecular identification of blowflies collected from Huachiachi.Since blowflies are holometabolous with morphologically different immature stages,the difficulties in the identification of larval stages limit their use as forensic indicators unless there is an alternative.Molecular taxonomy with its modern advances provides that alternative.In this context,molecular sequencing for forensic identification is performed here for the first time for blowflies of Huachiachi,Hangzhou. Five gene segments were specially selected for this purpose from the mitochondrial DNA (mt DNA =COI,CYTB,ND5) and ribosomal DNA(rDNA = ITS1 & ITS2) based on their popular use in forensic genetics.The 75 sequences yielded as a result were submitted to Genbank.We compared inter-specific variation at five gene loci for about 28 pairs of blowfly species(and also blowfly,housefly pairs).Among blowflies,(1) intra-specific sequence variation for ITS1 was in a reasonable range,i.e.0.29-1.16%.On the contrary inter-specific variation was quite high 7.8-57.3%;thus revealing its utility in congeneric species identifications.(2) ITS2 showed an inter-specific variation of 1.2-19%which illustrates a powerful resolution at species level yet at the same time shows an inevitable possibility of wrongful identification for sister species.(3) COI with an inter-specific divergence of 2.2-10.4%also indicated the likelihood of wrongful sister species identification.(4) On the other hand CYTB and ND5 regions with inter-specific divergence at 5.2-15.4%and 4.5-12.4%respectively showed a good resolution power for species level identifications.Among blowflies and housefly pairs(1) ITS2 proved as the most important gene marker for the family level differentiations with 30-35.2%variation.(2) COI with a variation of 10.3-13.5%showed no distinct demarcation between the "blowfly and housefly" pairs and some "blowfly and blowfly" pairs(10.4%).(3) CYTB with an inter-family divergence of 12.5-16.7%also showed no clear separation of species at the family level.(4) However ND5 provided distinct family level resolutions(16-17%).(5) Our results additionally demonstrated the usefulness of a group of concatenated gene segments,i.e.CO1,CYTB,ND5 and ITS2,in blowfly identifications based on phylogeny. (6) We also calculated an evolutionary timeline of blowflies based on our molecular data, previously published GenBank data,contemporary fossil evidence and theoretical assumptions.
We conclude that ITS1 as a novel sequence with low intra-and high inter-specific variability shows potential in identification of sister species that are quite similar at other gene loci.ND5 shows potential in identification of geographic populations of blowflies. ITS2 and ND5 both show family level resolution for flies of forensic importance.COI and ITS2 depict the evolutionary break for Calliphoridae and Muscidae at about 20-50 MYA.
In a parallel study,which is currently the first of its kind in the world,conducted in Pakistan,we explored insect fauna associated with the festival of Eid ul Azha(Eid).Thus, during the Eid time frame in 2008,in a remote Pakistani village,we examined(a) postmortem insect attractions on various sacrificial products,i.e.blood pools,meat, hides/skins,bones and offal(23 animals),and(b) and insect succession on 11 goat offal at five dump sites.Our results show:(1) Thirty five insect species were collected from 14 sites.(2) Carrion present outdoor attracted a diverse array of first wave of flies both by taxa and in numbers as compared to indoors where the initial wave includes only synanthrops. (3) During the present study we also carried out molecular seqauencing of blowflies for the first time in Pakistan.We sequenced three gene segments(COⅠ,COⅡand ITS2) for the Eid population of blowfly Ch.rufifaciesfrom D.I.Khan city of Pakistan.Additionally we sequenced same gene segments for the Eid population of Ch.rufifacies from Rawalpindi city of Pakistan.Massive transport of animals during the festival time throughout the country might facilitate invasion of forensically important fly species.So far our molecular sequence data for Ch.rufifacies population shows no significant variation for two different cities thus supporting our initial assumption that the Eid festival facilitates geneflow between different regions.Further forensic implications of these results in a broader context of Eid festval are discussed.
本论文分别调查分析了中国和巴基斯坦两国代表性区域法医昆虫的种类与种群演替规律,以及巴基斯坦传统节日—宰牲节对法医昆虫种类等的影响,并对杭州华家池重要类群丽蝇的分子鉴定技术进行了研究。
中国杭州华家池周围的蝇类(双翅目:Diptera)调查结果表明:(1)华家池共有蝇类法医昆虫12种,其中丽蝇科Calliphoridae7种,分别为大头金蝇Chrysomyamegacephala(Fabricius)、肥躯金蝇Ch.pinguis(Walker)、巨尾阿丽蝇Aldrichinagrahami(Aldrich)、丝光绿蝇Lucilia sericata(Meigen)、紫绿蝇L.porphyrina(Walker)、亮绿蝇L.illustris(Meigen)和瘦叶带绿蝇Hemipyrellia,igurriens(Wiedemann);麻蝇科Sarcophagidae一种,即肥须亚麻蝇Parasarcophagacrassipalpis(Macquart);蝇科Muscidae 4种,即家蝇Musca domestica(Linnaeus)、Musca sp.、Fannia sp.、Ophyra sp.;(2)大头金蝇是唯一全年都可以诱集到的种群,而且该种虫口数量占绝对的优势,而紫绿蝇虫口数量最小;(3)2007年1月至2008年12月的调查结果显示,丽蝇科7个种类、肥须亚麻蝇和家蝇的虫口数量在冬季均明显偏低,并且一月份只诱集到了大头金蝇和家蝇两种,二月份诱集到大头金蝇、肥躯金蝇、巨尾阿丽蝇、家蝇等四种,三月份诱集到了大头金蝇、肥躯金蝇、巨尾阿丽蝇、丝光绿蝇、亮绿蝇、肥须亚麻蝇、家蝇等7种;同时,统计分析结果表明虫口数量在这些月份之间没有显著差异,但冬夏种群虫口数量差异显著。
鉴于丽蝇幼期种类在形态上难以区分,本研究以华家池采集的丽蝇为材料,选用来自线粒体基因组的COⅠ、CYTB和ND5以及核糖体基因组的ITS1和ITS2共5个基因片段做分子标记,对丽蝇种类的分子鉴定进行了探索性研究。从7种丽蝇17个标本中扩增得到了75条基因序列并提交到GenBank。结果表明COⅠ、Cytb、ND5和ITS2可作为丽蝇种间分子鉴定的分予标记。ITS1序列可以作为丽蝇同属分子鉴定的分子标记。同时,研究比较了28对蝇(包括丽蝇、家蝇)5个基因位点的种间差异。结果表明:在丽蝇中,(1)ITS1的种内序列差异度为0.29-1.16%,而在种间水平上差异显著,为7.8-57.3%,表明ITS1能够用来进行同属内近缘种类鉴定;(2)ITS2的种间差异度为1.2-19%,表明该序列能够用于种的鉴别,但在姊妹中则可能出错;(3)COI序列的种内差异度在2.2-10.4%之间,也显示在姊妹种的鉴定上可能会错;(4)另一方面,CYTB和ND5序列种间分歧度分别为5.2-15.4%,4.5-12.4%,表明这两个序列均可成功用于种的鉴定。在丽蝇和家蝇组合中,(1)ITS2有30-35.2%的科间序列差异度,从而证明可作为最重要的基因标记用于科级水平的鉴定;(2)COⅠ的序列差异度10.3-13.5%表明在丽蝇科、家蝇科和某些丽蝇科内种类之间(10.4%)不能明显地区分;(3)CYTB序列的科间差异度12.5-16.7%也说明该序列在科技水平上不能明显区分;(4)但ND5序列的科间差异度16-17%则说明在可用于科级水平的鉴定;(5)结果还证明CO1、CYTB、ND5和ITS2等基因在鉴别丽蝇的系统发育关系中也很有用;(6)基于自己获得的基因序列以及GenBank中获得的分子数据,还推算了丽蝇科种类的演化关系。总之,ITS1序列具有低的种内和高的种间差异度,可望用于近缘种的区分;ND5可用于丽蝇不同地理种群的鉴别;ITS2和ND5均表现出科级水平的差异显著,COⅠ和ITS2序列分析表明,丽蝇科和家蝇科大约于20-50 MYA形成不同的进化分支。
在巴基斯坦穆斯林传统节日—宰牲节期间,大量动物尸体残留物抛弃在外,从而引起尸食类昆虫的大量繁殖,而目前尚无脊椎动物在户外垃圾场的降解以及与疾病传播方面的研究报道。本文以巴基斯坦一个典型乡镇为代表,2008年首次对宰牲节期间产生的血坑、肉、皮毛和骨头等14处场所进行了调查,共发现35种昆虫,其中腐肉上面发现的种类和数量最多;首次分析研究了分别来自巴基斯坦D.I.Khan城和Rawalpindi城宰牲节期间红颜金蝇Ch.rufifacies种群的COⅠ、COⅡ和ITS2基因序列,并加以对比分析,发现两个城市不同个体的红颜金蝇的分子序列分析完全相同。同时,在宰牲节期间的调查也发现牲畜交易过程提高了蝇蛆病的传播效率,两个城市的红颜金蝇不同个体分子序列分析没有差异的结果暗示了大量的动物贸易可能会携带并传播一些重要的蝇类,从而导致蝇类种群扩散。另外,还讨论了宰牲节对法医鉴定产生的深远影响。
The present study was aimed at achieving three main goals,to define the urban profile of blowflies,to devise a multiple gene approach for their molecular identification and to examine the association of blowflies and other carrion breeding insects with the Muslim "Festival of sacrifice i.e.Eid-ul-Azha".
Community structure of blowflies was determined in the urban settings of Huajiachi, Hangzhou,eastern China.Standard liver-sodium sulfide baits were placed along various urban facilities,including open sewerage drain,garbage dump sites,mulberry plantation, college building,pig farm and meat-vegetable markets.(1) We collected 12 species in all with nine species of major forensic importance including Chrysomya megacephala,Ch. pinguis,Aldrichina grahami,Lucilia sericata,L.illustris,L.porphyrina,Hemipyrellia ligurriens(Diptera:Calliphoridae),Musca domestica(Muscidae) and Parasarcophaga crassipalpis(Sarcophagidae).(2) ANOVA test statistics revealed significant mean population variation between species(F= 251.670) and months(F= 164.231).(3) Ch. megacephala was the only species observed throughout the study year.This species was the most dominant and abundant group with a population mean of 33±3.919 while L. porphyrina with a mean of 0.27±0.122 was the least represented species.(4) A special assemblage of seven species excluding the blowfly Ch.megacephala and housefly M. domestica were found absent in the January of two consecutive years,i.e.2007 and 2008. In February five species L.sericata,L.porphyrina,L.illustris,H.ligurriens and the flesh fly P.crassipalpis were absent in 2007-08 while in March only L.porphyrina and H. ligurriens were absent in 2007-08.Therefore,winter population means of all nine species were considerably low;January=3.3±0.246,February=1.14±0.472,December=0.25±0.76 and LSD analysis showed no significant difference for these months.However, statistical variation was significant when compared for winter and summer months.
The second phase of our study dealt with the molecular identification of blowflies collected from Huachiachi.Since blowflies are holometabolous with morphologically different immature stages,the difficulties in the identification of larval stages limit their use as forensic indicators unless there is an alternative.Molecular taxonomy with its modern advances provides that alternative.In this context,molecular sequencing for forensic identification is performed here for the first time for blowflies of Huachiachi,Hangzhou. Five gene segments were specially selected for this purpose from the mitochondrial DNA (mt DNA =COI,CYTB,ND5) and ribosomal DNA(rDNA = ITS1 & ITS2) based on their popular use in forensic genetics.The 75 sequences yielded as a result were submitted to Genbank.We compared inter-specific variation at five gene loci for about 28 pairs of blowfly species(and also blowfly,housefly pairs).Among blowflies,(1) intra-specific sequence variation for ITS1 was in a reasonable range,i.e.0.29-1.16%.On the contrary inter-specific variation was quite high 7.8-57.3%;thus revealing its utility in congeneric species identifications.(2) ITS2 showed an inter-specific variation of 1.2-19%which illustrates a powerful resolution at species level yet at the same time shows an inevitable possibility of wrongful identification for sister species.(3) COI with an inter-specific divergence of 2.2-10.4%also indicated the likelihood of wrongful sister species identification.(4) On the other hand CYTB and ND5 regions with inter-specific divergence at 5.2-15.4%and 4.5-12.4%respectively showed a good resolution power for species level identifications.Among blowflies and housefly pairs(1) ITS2 proved as the most important gene marker for the family level differentiations with 30-35.2%variation.(2) COI with a variation of 10.3-13.5%showed no distinct demarcation between the "blowfly and housefly" pairs and some "blowfly and blowfly" pairs(10.4%).(3) CYTB with an inter-family divergence of 12.5-16.7%also showed no clear separation of species at the family level.(4) However ND5 provided distinct family level resolutions(16-17%).(5) Our results additionally demonstrated the usefulness of a group of concatenated gene segments,i.e.CO1,CYTB,ND5 and ITS2,in blowfly identifications based on phylogeny. (6) We also calculated an evolutionary timeline of blowflies based on our molecular data, previously published GenBank data,contemporary fossil evidence and theoretical assumptions.
We conclude that ITS1 as a novel sequence with low intra-and high inter-specific variability shows potential in identification of sister species that are quite similar at other gene loci.ND5 shows potential in identification of geographic populations of blowflies. ITS2 and ND5 both show family level resolution for flies of forensic importance.COI and ITS2 depict the evolutionary break for Calliphoridae and Muscidae at about 20-50 MYA.
In a parallel study,which is currently the first of its kind in the world,conducted in Pakistan,we explored insect fauna associated with the festival of Eid ul Azha(Eid).Thus, during the Eid time frame in 2008,in a remote Pakistani village,we examined(a) postmortem insect attractions on various sacrificial products,i.e.blood pools,meat, hides/skins,bones and offal(23 animals),and(b) and insect succession on 11 goat offal at five dump sites.Our results show:(1) Thirty five insect species were collected from 14 sites.(2) Carrion present outdoor attracted a diverse array of first wave of flies both by taxa and in numbers as compared to indoors where the initial wave includes only synanthrops. (3) During the present study we also carried out molecular seqauencing of blowflies for the first time in Pakistan.We sequenced three gene segments(COⅠ,COⅡand ITS2) for the Eid population of blowfly Ch.rufifaciesfrom D.I.Khan city of Pakistan.Additionally we sequenced same gene segments for the Eid population of Ch.rufifacies from Rawalpindi city of Pakistan.Massive transport of animals during the festival time throughout the country might facilitate invasion of forensically important fly species.So far our molecular sequence data for Ch.rufifacies population shows no significant variation for two different cities thus supporting our initial assumption that the Eid festival facilitates geneflow between different regions.Further forensic implications of these results in a broader context of Eid festval are discussed.
引文
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