伞滑刃属线虫生物多样性与生物地理学
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摘要
生物地理学是以地球表层的生物群为研究对象,揭示生物多样性的空间分布规律,描述地球上生物数量和物种的各种分布形式的一门科学。结合植物检疫的工作实际,本课题研究论述了生物地理学在植物检疫中的作用,阐述了生物地理学理论在植物检疫和有害生物风险分析中的应用前景。植物检疫主要对象是外来有害生物,有害生物风险分析所涉及的有害生物的地理分布特征,与地理分布相关的生态因子(如气候、寄主和天敌),与扩散相关的非生态因子(如有害生物的生物学特性、传播和扩散途径),这些因子有许多是属于生物地理学的研究范畴。本课题研究对伞滑刃属线虫Bursaphelenchus Fuchs,1937的生物多样性和生物地理学特征进行了探讨,主要成果和创新点如下:
一、探讨了伞滑刃属线虫的生物多样性(Biodiversity),包括物种多样性和生态多样性。
在伞滑刃属线虫物种多样性(speciece diversity)研究方面,结合进出境木包装检疫和松材线虫疫情监测,开展了伞滑刃线虫种类鉴定和分类。从我国马尾松Pinus massoniana上发现伞滑刃属1个新种:拟小松伞滑刃线虫Bursaphelenchus parapinasteri sp. nov.,从进出境木包装板材和我国马尾松、湿地松P. elliotii上鉴定和报道了5个中国新记录种:伪伞滑刃线虫B. fraudulentus、食菌伞滑刃线虫B. fungivorus、树皮象伞滑刃线虫B. hylobianum、莱奴尔夫伞滑刃线虫B. rainulfi、泰国伞滑刃线虫B. thailandae、食菌伞滑刃线虫B. fungivorus。
系统收集和整理了至2007年为止的87个种。在此基础上编写出伞滑刃属87个种的物种名录;依据伞滑刃属线虫雄虫交合刺、尾部乳突、交合伞的形态,雌虫阴门及其阴门盖、后阴子宫囊和尾部形态、个体的总体形态特征等,编制了伞滑刃属87个种的检索表;依据交合刺形态特征将87种伞滑刃线虫按B. aberrans组、B. borealis组、B. eidmanni组、B. hunti组、B. piniperdae组和B. xylophilus组6个组重新进行分组。并比较全面地揭示伞滑刃属线虫的物种多样性,为伞滑刃属线虫种类鉴定和物种多样性研究提供新的依据。
在伞滑刃属线虫生态多样性(ecodiversity)研究方面,以伞滑刃属线虫各物种的寄主范围、传播媒介和传播途径、营养方式、生态位为依据,将87种伞滑刃线虫划分成4个生活类型:Ⅰ型为食真菌或自由生活的伞滑刃线虫,有22种;Ⅱ型为昆虫寄生型线虫,有8种;Ⅲ型为以昆虫作为传播媒介,食真菌的兼性寄生线虫,有55种;Ⅳ为型植物病原线虫,有2种。伞滑刃属线虫生活类型反映出该类线虫沿袭了滑刃目线虫从食真菌和低等丝状藻类线虫,向昆虫寄生线虫和植物寄生线虫演变的生态进化过程。
二、提出了伞滑刃属线虫地理分布与区系区划
根据收集的资料,揭示了世界不同地理区域的伞滑刃线虫物种丰富度与生态环境的关系.地理物种丰富度依次为:欧洲(42种)、亚洲(30种)、北美洲(21种)、加勒比海和中、南美洲(4种);对世界上不同国家和地区中含有伞滑刃线虫的种类进行归纳,应用GIS软件,将每个国家作为分布点,以种类及其种数数据作为每个分布点的属性值,叠加至软件提供的数字世界地图上,制作成伞滑刃属线虫种类地理分布的世界地图。这种地理分布图可以直观地表示世界上各个国家目前存在或发现的伞滑刃线虫种类和种数,表示出各种伞滑刃线虫目前在世界上的地理分布状况、传播范围;可以利用这种分布图根据地理位置和地理生态条件来分析各种伞滑刃线虫大体上的生存环境,为有害生物风险分析和植物检疫提供参考依据。
研究了伞滑刃属线虫在世界动物地理区的分布。根据地带性原则,以伞滑刃线虫属在各动物地理区及其邻近地区分布现状为依据,将该属线虫种类区分为广布种、共有种、特有种。探讨了伞滑刃线虫物种起源、自然扩散的途径以及人为扩散对物种分布的影响。
三、揭示了伞滑刃属线虫地理演变规律
伞滑刃属在世界起源和扩散过程中的地理分化体现了伞滑刃属线虫从食真菌和低等丝状藻类线虫,向昆虫寄生线虫和植物寄生线虫,从低级到高级的生态进化过程,表明伞滑刃属线虫的地理扩散与伞滑刃属线虫进化同时进行,伞滑刃属线虫在地理扩散中不断进化,在进化中逐渐扩散。研究表明伞滑刃线虫物种起源与生态位关系密切,在松科植物中伞滑刃属线虫具有丰富的物种。通过分析指出:物种在起源地与生态环境建立的和谐关系,是维持生态平衡的基础。松材线虫作为北美物种,与北美松树树种建立了稳定的平衡关系,对于北美松树树种危害性较小;一旦传入新区这种平衡被打破,松材线虫成为外来有害生物,表现极强的扩张性对新区松树破坏性极大。
四、结合中国松材线虫病的防控对松材线虫生物地理学进行了研究
嗜木伞滑刃线虫Bursaphelenchus xylophilus(Steiner & Buhrer,1934)Nickle,1970引起松树萎蔫病,俗称为松材线虫(pine wood nematode)或松树萎蔫线虫(pine wilt nematode),我国和世界上许多国家都将该线虫列为检疫对象。通过资料分析,阐述了松材线虫的世界分布、生物地理学特性、危害特点与检疫重要性;编制了松材线虫的世界分布地图。
根据松材线虫历年在我国的发生情况和对外公布的松材线虫分布状况,结合中国的地理和地形特点,运用GPS和GIS技术,研制了中国松材线虫疫区区域分布地图和中国松材线虫疫区扩散与地形地貌关系地图;根据研究资料制订了松材线虫分布危害的程度海拔高度区划指标:松材线虫分布低于海拔400m为松树萎蔫病严重发生区,海拔400-700m为中度发生区,海拔700-1000m为偶生区,海拔1000m以上为不发生区。划分了我国松材线虫4种地理理论发生区:①重度发生区、②轻度发生区、③偶发区、④无害区。确定长江三角洲、珠江三角洲和台湾岛作为我国松材线虫病3大易感中心。以上松材线虫的生物地理学研究结果,为我国松材线虫的检疫和、监控和预测提供了参考依据。
Biogeography is the scientific study of the geographic distribution of organisms. It aims to study the earth surface layer biota, reveals the spatial distribution of biodiversity laws and describes the biological quantity and species distribution pattern on the earth. In this dissertation, the biogeography and the plant quarantine was unified. The role of the biogeography in the plant quarantine, the application of biogeography in plant quarantine and pest risk analysis were expounded and discussed.
The main objects of the plant quarantine focus on invasive alien pests and the characteristics of geographical distribution of pests. The geographic distribution related to ecological factors (such as climate, host plants and natural enemies) and the proliferation of non-ecological factors (such as pest biology characteristic, transmission and dispersal route). Thus, many such factors are belong to the field of biogeography. Biodiversity and biogeography characters of the genus Bursaphelenchus Fuchs, 1937 were discussed. The main work and innovations in this dissertation are as follows:
1. Species diversity and ecological diversity of the genus Bursaphelenchus were elaborated. The taxonomy of Bursaphelenchus spp. was identified during the quarantine of entry-exit wood packaging and the epidemic monitoring process of pin wood nematode. A new species was identified as B. parapinasteri sp. nov. from masson pine (Pinus massoniana). B. fraudulentus, B. fungivorus, B. hylobianum, B. rainulfi, and B. thailandea were reported as new records in China.
The identification key to 87 species of Bursaphelenchus spp. was established based on the characteristics of spicule, tail papilla, shape of copulatory bursa of male, vulva epiptygma, postuterine branch, tail shape of female, the whole morphology characteristics and so on. According to morphological characteristics of spicule, the genus Bursaphelenchus was re-individed into 6 groups : B. aberrans group, B. borealis group, B. eidmanni group, B. hunti group, B. piniperdae group and B. xylophilus group. All those results above are based on relevant domestic and international references which were systematically collected, sorted and processed up to 2007 about this genus.
In eco-diversity, 87 species of the genus Bursaphelenchus were individed into 4 life-styles according to their host range, transmission, nutrition support methods and niche. Life-styleⅠ: 22 species of them were considered as fungal-feeding or free-living Bursaphelenchus spp; Life-styleⅡ:8 species were defined as entomogenous nematodes; Life-styleⅢ: There are 55 members transmitting by insect vectors,fungal-feeding or facultative parasite nematodes; Life-styleⅣ:2 species are plant pathogenic nematodes.
Life styles reflect a certain relationship of inheritance in ecologically evolutional process between Bursaphelenchus spp. and Aphelenchida nematodes from fungal-feeding and filamentous algae-feeding to entomogenous nematodes and plant pathogenic nematodes.
2. Geographical distribution and fauna of the genus Bursaphelenchus in the world were discussed.
Based on the geographic distribution of Bursaphelenchus spp., the richness of the genus in different geographic regions and the relationship between species richness and the ecological environment were revealed. Geography species richness are 42 species in Europe, 30 species in Asia, 21 species in North America, 4 species in the Caribbean and Central and South America. According to the relevant domestic and international references, species of Bursaphelenchus spp. of different countries and regions were summarized, countries as distribution spot, species and the quantity of species for attribute value and superimposes on digit world map which provides by GIS software , the world geographical distribution map of Bursaphelenchus spp. was provided.
It is intuitive that geographic distribution of the world that all countries are currently exist or found species, expressed species of Bursaphelenchus spp. present in the world, the geographical distribution of the communication sphere. According to ecological distribution and geographic location of the species, the survival environment of Bursaphelenchus spp. and pest risks were provided to plant quarantine. Distribution of zoogeographical regions of the world of Bursaphelenchus spp. was performed.
Based on the zonality principle and distribution status of zoogeographical regions and areas adjacent to the status of Bursaphelenchus spp. has been divided into cosmopolitan species, mutual species and endemic species., mutual species and endemic species. The effect of natural dispersal route and artificial dispersal route of Bursaphelenchus spp. was discussed.
3. The geographic dispersion law of the genus Bursaphelenchus was summarized.
Geographical differentiation of the genus Bursaphelenchus in its origin and dispersion process, reflects the evolution process from fungal-feeding and filamentous algae-feeding to entomogenous nematodes and plant pathogenic nematodes from the preliminary class to high class. It indicted the parallel relationship between geographical extension and species evolution. The species evolution is often contemporaneous with the geographical dispersion.
The results also indicated that there was a close relationship between the species origin and ecological niche of Bursaphelenchus spp. Our study showed that the harmonious relationship between species origin location and ecological environment sustains the ecological balance. The Bursaphelenchus spp. as the species in North America has been formed a stable equilibrium to the North America pine with lower threat to North American pine. Once the balance is disrupted, the nematodes as the alien pests enormously spread to new area of pine trees and therefore resulted in considerable destruction.
4. The biogeography of B. xylophilus was studied with the disease control in China.
Bursaphelenchus xylophilus (Steiner & Buhrer, 1934) Nickle, 1970, was known as a pine wood nematode or pine wilt nematode (PWN), which was the quarantine pest in China and some other countries. The biogeographic characteristics, damage characteristic and quarantine significance of PWN were elaborated. The distribution map was established based on available data.
According to the occurrence status of Bursaphelencesus spp. in China, publishing data of distribution status of PWN and the Chinese geography and topographic characteristics, the distribution regional map and potential invasion area in China were planed by using GPS and GIS technology. Thus, the damage degree by PWN was designed as the serious infection ranges (<400 m elevation), medium infection ranges (400-700 m elevation) and incidental occurrence ranges (700-1000 m) in light of altitude.
Theoretical geographical occurrence area has individed into 4 types :①severe occurred area,②mild occurred area,③incidental occurred area,④safety area. Yangtze River Delta, Pearl River Delta and Taiwan island are consided as pine wilt disease susceptibility centre in China. The above results of biogeography provided a reference for PWN quarantine, monitoring and forecast in China.
一、探讨了伞滑刃属线虫的生物多样性(Biodiversity),包括物种多样性和生态多样性。
在伞滑刃属线虫物种多样性(speciece diversity)研究方面,结合进出境木包装检疫和松材线虫疫情监测,开展了伞滑刃线虫种类鉴定和分类。从我国马尾松Pinus massoniana上发现伞滑刃属1个新种:拟小松伞滑刃线虫Bursaphelenchus parapinasteri sp. nov.,从进出境木包装板材和我国马尾松、湿地松P. elliotii上鉴定和报道了5个中国新记录种:伪伞滑刃线虫B. fraudulentus、食菌伞滑刃线虫B. fungivorus、树皮象伞滑刃线虫B. hylobianum、莱奴尔夫伞滑刃线虫B. rainulfi、泰国伞滑刃线虫B. thailandae、食菌伞滑刃线虫B. fungivorus。
系统收集和整理了至2007年为止的87个种。在此基础上编写出伞滑刃属87个种的物种名录;依据伞滑刃属线虫雄虫交合刺、尾部乳突、交合伞的形态,雌虫阴门及其阴门盖、后阴子宫囊和尾部形态、个体的总体形态特征等,编制了伞滑刃属87个种的检索表;依据交合刺形态特征将87种伞滑刃线虫按B. aberrans组、B. borealis组、B. eidmanni组、B. hunti组、B. piniperdae组和B. xylophilus组6个组重新进行分组。并比较全面地揭示伞滑刃属线虫的物种多样性,为伞滑刃属线虫种类鉴定和物种多样性研究提供新的依据。
在伞滑刃属线虫生态多样性(ecodiversity)研究方面,以伞滑刃属线虫各物种的寄主范围、传播媒介和传播途径、营养方式、生态位为依据,将87种伞滑刃线虫划分成4个生活类型:Ⅰ型为食真菌或自由生活的伞滑刃线虫,有22种;Ⅱ型为昆虫寄生型线虫,有8种;Ⅲ型为以昆虫作为传播媒介,食真菌的兼性寄生线虫,有55种;Ⅳ为型植物病原线虫,有2种。伞滑刃属线虫生活类型反映出该类线虫沿袭了滑刃目线虫从食真菌和低等丝状藻类线虫,向昆虫寄生线虫和植物寄生线虫演变的生态进化过程。
二、提出了伞滑刃属线虫地理分布与区系区划
根据收集的资料,揭示了世界不同地理区域的伞滑刃线虫物种丰富度与生态环境的关系.地理物种丰富度依次为:欧洲(42种)、亚洲(30种)、北美洲(21种)、加勒比海和中、南美洲(4种);对世界上不同国家和地区中含有伞滑刃线虫的种类进行归纳,应用GIS软件,将每个国家作为分布点,以种类及其种数数据作为每个分布点的属性值,叠加至软件提供的数字世界地图上,制作成伞滑刃属线虫种类地理分布的世界地图。这种地理分布图可以直观地表示世界上各个国家目前存在或发现的伞滑刃线虫种类和种数,表示出各种伞滑刃线虫目前在世界上的地理分布状况、传播范围;可以利用这种分布图根据地理位置和地理生态条件来分析各种伞滑刃线虫大体上的生存环境,为有害生物风险分析和植物检疫提供参考依据。
研究了伞滑刃属线虫在世界动物地理区的分布。根据地带性原则,以伞滑刃线虫属在各动物地理区及其邻近地区分布现状为依据,将该属线虫种类区分为广布种、共有种、特有种。探讨了伞滑刃线虫物种起源、自然扩散的途径以及人为扩散对物种分布的影响。
三、揭示了伞滑刃属线虫地理演变规律
伞滑刃属在世界起源和扩散过程中的地理分化体现了伞滑刃属线虫从食真菌和低等丝状藻类线虫,向昆虫寄生线虫和植物寄生线虫,从低级到高级的生态进化过程,表明伞滑刃属线虫的地理扩散与伞滑刃属线虫进化同时进行,伞滑刃属线虫在地理扩散中不断进化,在进化中逐渐扩散。研究表明伞滑刃线虫物种起源与生态位关系密切,在松科植物中伞滑刃属线虫具有丰富的物种。通过分析指出:物种在起源地与生态环境建立的和谐关系,是维持生态平衡的基础。松材线虫作为北美物种,与北美松树树种建立了稳定的平衡关系,对于北美松树树种危害性较小;一旦传入新区这种平衡被打破,松材线虫成为外来有害生物,表现极强的扩张性对新区松树破坏性极大。
四、结合中国松材线虫病的防控对松材线虫生物地理学进行了研究
嗜木伞滑刃线虫Bursaphelenchus xylophilus(Steiner & Buhrer,1934)Nickle,1970引起松树萎蔫病,俗称为松材线虫(pine wood nematode)或松树萎蔫线虫(pine wilt nematode),我国和世界上许多国家都将该线虫列为检疫对象。通过资料分析,阐述了松材线虫的世界分布、生物地理学特性、危害特点与检疫重要性;编制了松材线虫的世界分布地图。
根据松材线虫历年在我国的发生情况和对外公布的松材线虫分布状况,结合中国的地理和地形特点,运用GPS和GIS技术,研制了中国松材线虫疫区区域分布地图和中国松材线虫疫区扩散与地形地貌关系地图;根据研究资料制订了松材线虫分布危害的程度海拔高度区划指标:松材线虫分布低于海拔400m为松树萎蔫病严重发生区,海拔400-700m为中度发生区,海拔700-1000m为偶生区,海拔1000m以上为不发生区。划分了我国松材线虫4种地理理论发生区:①重度发生区、②轻度发生区、③偶发区、④无害区。确定长江三角洲、珠江三角洲和台湾岛作为我国松材线虫病3大易感中心。以上松材线虫的生物地理学研究结果,为我国松材线虫的检疫和、监控和预测提供了参考依据。
Biogeography is the scientific study of the geographic distribution of organisms. It aims to study the earth surface layer biota, reveals the spatial distribution of biodiversity laws and describes the biological quantity and species distribution pattern on the earth. In this dissertation, the biogeography and the plant quarantine was unified. The role of the biogeography in the plant quarantine, the application of biogeography in plant quarantine and pest risk analysis were expounded and discussed.
The main objects of the plant quarantine focus on invasive alien pests and the characteristics of geographical distribution of pests. The geographic distribution related to ecological factors (such as climate, host plants and natural enemies) and the proliferation of non-ecological factors (such as pest biology characteristic, transmission and dispersal route). Thus, many such factors are belong to the field of biogeography. Biodiversity and biogeography characters of the genus Bursaphelenchus Fuchs, 1937 were discussed. The main work and innovations in this dissertation are as follows:
1. Species diversity and ecological diversity of the genus Bursaphelenchus were elaborated. The taxonomy of Bursaphelenchus spp. was identified during the quarantine of entry-exit wood packaging and the epidemic monitoring process of pin wood nematode. A new species was identified as B. parapinasteri sp. nov. from masson pine (Pinus massoniana). B. fraudulentus, B. fungivorus, B. hylobianum, B. rainulfi, and B. thailandea were reported as new records in China.
The identification key to 87 species of Bursaphelenchus spp. was established based on the characteristics of spicule, tail papilla, shape of copulatory bursa of male, vulva epiptygma, postuterine branch, tail shape of female, the whole morphology characteristics and so on. According to morphological characteristics of spicule, the genus Bursaphelenchus was re-individed into 6 groups : B. aberrans group, B. borealis group, B. eidmanni group, B. hunti group, B. piniperdae group and B. xylophilus group. All those results above are based on relevant domestic and international references which were systematically collected, sorted and processed up to 2007 about this genus.
In eco-diversity, 87 species of the genus Bursaphelenchus were individed into 4 life-styles according to their host range, transmission, nutrition support methods and niche. Life-styleⅠ: 22 species of them were considered as fungal-feeding or free-living Bursaphelenchus spp; Life-styleⅡ:8 species were defined as entomogenous nematodes; Life-styleⅢ: There are 55 members transmitting by insect vectors,fungal-feeding or facultative parasite nematodes; Life-styleⅣ:2 species are plant pathogenic nematodes.
Life styles reflect a certain relationship of inheritance in ecologically evolutional process between Bursaphelenchus spp. and Aphelenchida nematodes from fungal-feeding and filamentous algae-feeding to entomogenous nematodes and plant pathogenic nematodes.
2. Geographical distribution and fauna of the genus Bursaphelenchus in the world were discussed.
Based on the geographic distribution of Bursaphelenchus spp., the richness of the genus in different geographic regions and the relationship between species richness and the ecological environment were revealed. Geography species richness are 42 species in Europe, 30 species in Asia, 21 species in North America, 4 species in the Caribbean and Central and South America. According to the relevant domestic and international references, species of Bursaphelenchus spp. of different countries and regions were summarized, countries as distribution spot, species and the quantity of species for attribute value and superimposes on digit world map which provides by GIS software , the world geographical distribution map of Bursaphelenchus spp. was provided.
It is intuitive that geographic distribution of the world that all countries are currently exist or found species, expressed species of Bursaphelenchus spp. present in the world, the geographical distribution of the communication sphere. According to ecological distribution and geographic location of the species, the survival environment of Bursaphelenchus spp. and pest risks were provided to plant quarantine. Distribution of zoogeographical regions of the world of Bursaphelenchus spp. was performed.
Based on the zonality principle and distribution status of zoogeographical regions and areas adjacent to the status of Bursaphelenchus spp. has been divided into cosmopolitan species, mutual species and endemic species., mutual species and endemic species. The effect of natural dispersal route and artificial dispersal route of Bursaphelenchus spp. was discussed.
3. The geographic dispersion law of the genus Bursaphelenchus was summarized.
Geographical differentiation of the genus Bursaphelenchus in its origin and dispersion process, reflects the evolution process from fungal-feeding and filamentous algae-feeding to entomogenous nematodes and plant pathogenic nematodes from the preliminary class to high class. It indicted the parallel relationship between geographical extension and species evolution. The species evolution is often contemporaneous with the geographical dispersion.
The results also indicated that there was a close relationship between the species origin and ecological niche of Bursaphelenchus spp. Our study showed that the harmonious relationship between species origin location and ecological environment sustains the ecological balance. The Bursaphelenchus spp. as the species in North America has been formed a stable equilibrium to the North America pine with lower threat to North American pine. Once the balance is disrupted, the nematodes as the alien pests enormously spread to new area of pine trees and therefore resulted in considerable destruction.
4. The biogeography of B. xylophilus was studied with the disease control in China.
Bursaphelenchus xylophilus (Steiner & Buhrer, 1934) Nickle, 1970, was known as a pine wood nematode or pine wilt nematode (PWN), which was the quarantine pest in China and some other countries. The biogeographic characteristics, damage characteristic and quarantine significance of PWN were elaborated. The distribution map was established based on available data.
According to the occurrence status of Bursaphelencesus spp. in China, publishing data of distribution status of PWN and the Chinese geography and topographic characteristics, the distribution regional map and potential invasion area in China were planed by using GPS and GIS technology. Thus, the damage degree by PWN was designed as the serious infection ranges (<400 m elevation), medium infection ranges (400-700 m elevation) and incidental occurrence ranges (700-1000 m) in light of altitude.
Theoretical geographical occurrence area has individed into 4 types :①severe occurred area,②mild occurred area,③incidental occurred area,④safety area. Yangtze River Delta, Pearl River Delta and Taiwan island are consided as pine wilt disease susceptibility centre in China. The above results of biogeography provided a reference for PWN quarantine, monitoring and forecast in China.
引文
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