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有孔团水虱(Sphaeroma terebrans)属于甲壳纲(Crustacea)等足目(Isopoda)扇肢亚目(Flabellifera)团水虱科(Sphaeromatidae),团水虱属(Sphaeroma)。2013年以来,我国海南、广西、深圳等地陆续发生有孔团水虱造成红树林大面积死亡的事件,严重爆发可使红树林死亡面积以年均66%增长[1-2]。
目前国外团水虱主要集中在红茄冬(Rhizophora mucronata Poir.in Lam.)上,而我国遭受团水虱危害的主要树种是白骨壤(Avicennia marina (Forsk.) Vierh.)、木榄(Bruguiera gymnorhiza (L.) Lam.)、海莲(Bruguiera sexangula Lamk.)等红树种类。另外国外没有出现大面积红树林死亡的报道。在团水虱造成红树林严重危害之前,国内极少有文献报道,其生物学和防治的内容基本空白。关于有孔团水虱相关研究多数集中在其本身的生物学特征。结合我国各地的发生情况和前期的调查,整理和归纳了有孔团水虱的研究情况,以期对有孔团水虱的防治和综合治理有所帮助。
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在中国,与红树林密切相关的团水虱种类主要是有孔团水虱和光背团水虱(S. retrolaevis)两种,两者的名称常被混淆[1-2]。但这两者对红树林造成的影响差异大,从形态上正确区分非常必要。表1是两者的主要形态差异。
表 1 光背团水虱与有孔团水虱的主要区别
Table 1. The main difference between Sphaeroma terebrans and Sphaeroma retrolaevis
特征区别Distinctive features 光背团水虱Sphaeroma retrolaevis 有孔团水虱Sphaeroma terebrans 成年个体大小Adult size 可达10 mm Up to 10 mm 可达12 mm Up to 12 mm 第 5 ~7 胸节5th to 7th thoracic segments 无凸起No bulge 各具4个凸起 Each with 4 bulges 雄性最后1对足Male last pair of feet 未骨化Not ossification 骨化Ossification 下颌骨Mandible 接近Contiguity 分开Detachment 腹部末端形态Abdominal terminal morphology 平截Truncature 近抛物线 Near parabola 蛀孔形态Wormhole form 纵向 Parallel 横向 Cross -
国外主要分布在印度、斯里兰卡、泰国、印度尼西亚、澳大利亚、巴布亚新几内亚、美国、巴西等国家。国内分布在浙江以南沿海地区,其中海南、广州和广西三省内团水虱导致红树林大面积死亡,近期深圳也有致死情况发生。
物理障碍、地理隔离、海洋环流共同影响着有孔团水虱的种群地理分布结构[3-4]。有孔团水虱的区域分布可能受一些非生物因素的影响,包括盐度、水温、悬浮物、潮汐以及浸淹的影响[5-8]。
有孔团水虱生活在红树林根系所形成的潮间带生境中[6, 9],通常在红树植物的气生根中蛀孔居住。在肯尼亚岛屿和内陆空间尺度较大的红树林,位于低潮带尤其是位于潮沟边缘或向海边缘泥质或沙质基底的红茄冬,受有孔团水虱侵扰普遍且强度大,大部分根系被毁坏或死亡,而位于高潮带的红茄冬未发现其危害的迹象[10]。有孔团水虱危害红茄冬林支柱根的强度不一,仅在垂直高度1 m范围内,死亡根和受损根所占比例不等,但气生根受损一般发生在向海一侧的低水位泥质区。在肯尼亚和桑给巴尔岛沿岸红树林区,有孔团水虱是否危害红茄冬气生根一般取决于两个因素,即水位和沉积物类型。随着潮间带高度的增加,等足类甲壳动物浸在水中的时间减少,这不仅限制了它们的觅食时间,而且减少了其侵害根的时间。而泥质底质水体中的有机物要比沙质区的丰富,因而有孔团水虱在泥质区更易获取食物[11]。潮间带的基底沉积类型、水位高度及是否临水是影响有孔团水虱丰度的主要因素[10]。
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有孔团水虱属于甲壳动物等足类团水虱科,该科动物通常居住在热带或亚热带的红树植物的气生根中。一般来说,有孔团水虱的整个生命周期都在同一片的红树林中。Baratti等[12]对非洲和美国5个红树林生态系统的有孔团水虱进行采样分析,提供了不同地区种群之间遗传分化的明确证据:持续性低的种内变化和持续性高的种群间距离表明几乎所有的有孔团水虱线粒体的差异(细胞色素氧化酶I)体现在种群间而不是在种群内,其中平均单倍型多样性(h)是0.71%,平均核苷酸多样性(π)是0.34%;最小生成树(MST)揭示了一个复杂的模式:3个主要的单倍型类群对应的所研究地理位置分布在网络中。这表明了该种有一个悠久的进化史和种群间的基因流动受到了限制。考虑到类群相关特性,遗传分化程度并不能完全评价类群的分类地位,但在种群分支(A vs B+C=21%; A, B+C vs D=22%)的遗传分化可参考已有文献中描述的甲壳动物[12-13]。以甲壳动物来说,COI基因存在16%的差异可认定为两个物种[13]。因此,在属于不同类群种群间的核苷酸差异的百分率已经远大于同属不同物种间最小的极限[13]。Baratti等[12]认为有孔团水虱种群间大的遗传距离可以表明该分类群不是一个单一物种而是一个复杂的分类群,该类群的分类地位需重新定义。
为了了解有孔团水虱潜在的遗传分化和地理分布结构,研究人员对其开展系统地理学分析[14]。该物种在红树林根中度过其整个生命周期,具有较低的主动扩散能力且没有早期扩散阶段。由于这个物种分布于海洋,远程扩散机制(如通过浮木进行的漂流扩散)对其扩散分布是极其重要的。然而,研究人员在相邻的栖息地发现了高水平的遗传分化和低水平的基因流,与海岸地形相关的物理障碍可能有助于减少基因流。世界范围的有孔团水虱似乎不只包含一种种类。
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有孔团水虱无浮游幼体阶段,其整个生活史几乎全部在洞穴中度过,加之其天敌较少,这些导致有孔团水虱的有效防治途径较为匮乏。
国外关于团水虱防治的研究较少。Cragg团队提出化学处理和防腐剂会影响等足类动物的定殖[23]。Huynh等表明拟除虫菊酯类杀虫剂对等足类动物是有害的[54]。Roshaven发现,极端高温和极高盐度的单独或组合使用都可限制成年有孔团水虱的生存[55]。经铜铬砷或杂酚油表层涂层处理的木材,可以有效控制咀虫,在某些情况下也可以有效控制蛀木水虱和团水虱。团水虱的种群数量控制和健康的红树林群落可能是控制它们的关键。
我国防治研究相对较多。2013年以来,海南东寨港红树林保护区团水虱爆发成灾后,部分科研单位相继开展了红树林团水虱的防治研究工作,进行了有意义的技术探索并陆续申请了有关的发明专利(表2),积累了一定的防治经验。
表 2 国内关于红树林团水虱防治的相关专利申请
Table 2. Related patent application for the control of mangrove Sphaeroma sp. in China
专利发明人
The patent inventor专利名称
Patent专利简要说明
Brief description of patent何雪香,等[56]
He et al《防控团水虱的双重药剂及应用该药剂防控团水虱的方法》《The double agent for the control of and Sphaeroma sp. and the method for the control of the isopoda》 配制化学药剂进行涂抹Prepare chemicals to apply 刘文爱,等[57]
Liu et al《一种红树林团水虱虫害的防治方法》《The method for controlling Sphaeroma sp. of mangrove forest》 利用食盐进行涂抹或喷洒Apply or spray with salt 管伟,等[58]
Guan et al《一种快速防治团水虱的烟剂、烟熏装置和方法》《The invention relates to fumicant、a fumigating device and a method for rapid control of Sphaeroma sp.》 配制烟剂进行烟熏Smudging with The fumigant 吕晓波,等[59]
Lv et al《一种运用隔离原理治理红树林团水虱危害的物理防治方法》《A physical prevention method for using the principle of isolation to control the damage of mangrove Sphaeroma sp.》 使用塑料薄膜将团水虱物理隔离Use plastic film to physically separate the Sphaeroma sp. 黄勃,等[60]
Huang et al《防治团水虱和船蛆钻孔海洋生物的药物及其使用方法》《Medicine and method for controlling marine organisms Sphaeroma sp. and shipworm》 配制药剂进行涂抹,并涂上桐油隔离Applying medicaments to control and tung oil to segregation 刘文爱,等[61]
Liu et al《一种利用二氧化氯防治红树林团水虱虫害的方法》《A method to control mangrove Sphaeroma sp.by chlorine dioxide》 配制高浓度二氧化氯溶液进行喷洒Spraying high concentration chlorine dioxide to control 当前的团水虱防治专利主要集中在团水虱生物体的灭杀上,需要每木处理,不便操作、成本较高;防治技术手段不能从根本上解决团水虱的危害问题,对当地红树林生态环境也存在着一些不利的影响,因而需要继续研究探索更高效、环保的技术手段。
造成红树林团水虱灾害爆发原因是多方面的,通过研究寻找共性规律,揭示团水虱爆发的机理,为团水虱的科学治理提供科学基础;将团水虱监测纳入红树林生态监测体系中,鉴于其巨大的破坏作用,迫切需要研发红树林团水虱的灾害预警技术;重视团水虱灾后的红树林恢复重建,一方面需要通过人工造林恢复红树林的种群数量,另一方面是对大量受损退化红树植株的复状抚育。
团水虱个体较小,各种类的形态类似,体色容易受环境影响而有所不同,急需通过分子生物学的方法,通过基因测定加强有孔团水虱的鉴定,物种间的变异的研究。
有孔团水虱的爆发与红树林周边环境密切相关,指向性强,易于观测,是衡量红树林生态系统是否健康的重要指标。此外,由于团水虱链接了环境改变—红树林退化—红树林生态系统崩溃,因此在方法论上有孔团水虱是一个研究生态环境改变与自然生态系统演化关系的自然模板。
红树林顶级杀手—有孔团水虱的研究进展
A Top Killer of Mangroves – Research Progress of Sphaeroma terebrans
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摘要:
目的 我国多地多次发生有孔团水虱 (Sphaeroma terebrans) 造成红树林死亡的事件。目前国内关于该团水虱的研究较少,国外有较长的研究历史但是研究时间和地点比较分散,通过借鉴前人的研究成果,以方便我们了解并防控该虫。 方法 查询中外文献、专利,并结合本人已开展的研究,主要针对有孔团水虱的形态、分布和分类、生物学特征、天敌,对红树林的危害特征和红树林对其的响应,以及目前的防治手段等方面着手进行总结。 结果 有孔团水虱分布范围很广,其形态特征存在地理性差异,基因研究表明有孔团水虱种群间不是一个单一物种而是一个复杂的分类群。国外有孔团水虱主要蛀孔于红树林气生根,国内除了气生根外还蛀孔于红树植物基干,且前者鲜有致死红树林的报道。对于蛀孔的生物学意义争议很大,既有可引起的“生态灾难”,也有可给红树林带来的益处,但前者的支持者较多。团水虱一生至少蜕皮4次,两年可育3代。红树林气生根上团水虱空洞若数量较少,可自行愈合。有文献记录几种海洋生物可捕食团水虱,但天敌种类较少;国内已有关于团水虱的控制方法,但操作性差,且不能从根本上解决问题,仍需加强团水虱的防控相关研究,另外,开展团水虱灾害预警技术和灾后重建技术也迫在眉睫。 Abstract:Objective There have been many incidents in many places in China that have caused the death of mangroves by Sphaeroma terebrans. At present, there are few studies on the S. terebrans in China and there is a long research history abroad, but the research are relatively scattered. Method By researching Chinese and foreign literatures and patents, and combining with the research carried out, this paper focuses on the morphology, distribution and classification, biological characteristics, natural enemies of S. terebrans, and its damage to mangroves, the response of mangroves to S. terebrans, as well as the current prevention measures. Result The distribution range of S. terebrans is very wide, and there are geographical differences in its morphological characteristics. Genetic studies have shown that the population of S. terebrans is not a single species but a complex taxonomic group. In foreign countries, S. terebrans mainly bores holes in mangrove aerial roots, In China, they also bores holes in mangrove plant base in addition to aerial roots, and there are few reports of the former killing mangroves. The biological significance of borer holes is controversial. It can either cause ecological disaster or bring benefits to mangroves, but the former has more supporters. S. terebrans moults at least four times in a lifetime and can have three generations in two years. If the number of S. terebrans cavities on mangrove aerial roots is small, they can heal themselves. It has been documented that several marine organisms can prey on S. terebrans, but they have few kinds of natural enemies. There are some control methods for S. terebrans in China, but they are of poor operability and the problem cannot be solved fundamentally. Relevant research on prevention and control of S. terebrans still needs to be strengthened. In addition, early warning technology for S. terebrans disaster and post-disaster reconstruction technology are also imminent. -
Key words:
- Sphaeroma terebrans
- / mangrove
- / biological characteristics
- / prevention and control
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表 1 光背团水虱与有孔团水虱的主要区别
Table 1. The main difference between Sphaeroma terebrans and Sphaeroma retrolaevis
特征区别Distinctive features 光背团水虱Sphaeroma retrolaevis 有孔团水虱Sphaeroma terebrans 成年个体大小Adult size 可达10 mm Up to 10 mm 可达12 mm Up to 12 mm 第 5 ~7 胸节5th to 7th thoracic segments 无凸起No bulge 各具4个凸起 Each with 4 bulges 雄性最后1对足Male last pair of feet 未骨化Not ossification 骨化Ossification 下颌骨Mandible 接近Contiguity 分开Detachment 腹部末端形态Abdominal terminal morphology 平截Truncature 近抛物线 Near parabola 蛀孔形态Wormhole form 纵向 Parallel 横向 Cross 表 2 国内关于红树林团水虱防治的相关专利申请
Table 2. Related patent application for the control of mangrove Sphaeroma sp. in China
专利发明人
The patent inventor专利名称
Patent专利简要说明
Brief description of patent何雪香,等[56]
He et al《防控团水虱的双重药剂及应用该药剂防控团水虱的方法》《The double agent for the control of and Sphaeroma sp. and the method for the control of the isopoda》 配制化学药剂进行涂抹Prepare chemicals to apply 刘文爱,等[57]
Liu et al《一种红树林团水虱虫害的防治方法》《The method for controlling Sphaeroma sp. of mangrove forest》 利用食盐进行涂抹或喷洒Apply or spray with salt 管伟,等[58]
Guan et al《一种快速防治团水虱的烟剂、烟熏装置和方法》《The invention relates to fumicant、a fumigating device and a method for rapid control of Sphaeroma sp.》 配制烟剂进行烟熏Smudging with The fumigant 吕晓波,等[59]
Lv et al《一种运用隔离原理治理红树林团水虱危害的物理防治方法》《A physical prevention method for using the principle of isolation to control the damage of mangrove Sphaeroma sp.》 使用塑料薄膜将团水虱物理隔离Use plastic film to physically separate the Sphaeroma sp. 黄勃,等[60]
Huang et al《防治团水虱和船蛆钻孔海洋生物的药物及其使用方法》《Medicine and method for controlling marine organisms Sphaeroma sp. and shipworm》 配制药剂进行涂抹,并涂上桐油隔离Applying medicaments to control and tung oil to segregation 刘文爱,等[61]
Liu et al《一种利用二氧化氯防治红树林团水虱虫害的方法》《A method to control mangrove Sphaeroma sp.by chlorine dioxide》 配制高浓度二氧化氯溶液进行喷洒Spraying high concentration chlorine dioxide to control -
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