林业科学  2016, Vol. 52 Issue (2): 134-139   PDF    
DOI: 10.11707/j.1001-7488.20160217
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文章信息

王文婷, 肖洒, 黄河清, 李凯, 张东, 初红军, 国有清, 高万里
Wang Wengting, Xiao Sa, Huang Heqing, Li Kai, Zhang Dong, Chu Hongjun, Guo Youqing, Gao Wanli
我国蒙新区、青藏区马胃蝇多样性及感染分析
Diversity and Infection of Gasterophilus spp. in Mongol-Xinjiang Region and Qinghai Tibet Region
林业科学, 2016, 52(2): 134-139
Scientia Silvae Sinicae, 2016, 52(2): 134-139.
DOI: 10.11707/j.1001-7488.20160217

文章历史

收稿日期:2015-02-06
修回日期:2015-08-23

作者相关文章

王文婷
肖洒
黄河清
李凯
张东
初红军
国有清
高万里

我国蒙新区、青藏区马胃蝇多样性及感染分析
王文婷1, 肖洒1, 黄河清1, 李凯1, 张东1, 初红军2, 国有清3, 高万里4    
1. 北京林业大学自然保护区学院 北京 100083;
2. 新疆卡拉麦里山有蹄类自然保护区阿勒泰管理站 阿勒泰 836500;
3. 内蒙古赤峰市克什克腾旗青山林场 赤峰 025366;
4. 青海省林业厅 西宁 810008
摘要[目的] 研究新疆、内蒙、青海3个地区的草原、荒漠草原和高寒草原马胃蝇蛆病的发生情况、感染多样性和区系组成。[方法] 采用伊维菌素于同一年冬季分别对蒙新区的新疆卡拉麦里有蹄类自然保护区普氏野马、内蒙古多伦县当地家马以及青藏区的青海省玛多县当地家马进行驱虫,对获取的马胃蝇数量、种类进行调查和分析。[结果] 1)马胃蝇蛆病发生普遍,感染率在3个地区马科动物中均为100%,感染程度有明显的地域性:蒙新区重于青藏区;新疆卡拉麦里有蹄类自然保护区马胃蝇感染数量显著高于同区的内蒙古多伦县,后者的感染量明显高于青藏区的玛多县; 2)同属于蒙新区的新疆卡拉麦里有蹄类自然保护区和内蒙古多伦县具有相同的马胃蝇种类,所涉及的6个种(黑腹胃蝇、鼻胃蝇、肠胃蝇、黑角胃蝇、红尾胃蝇和裸节胃蝇)均为我国现有记录种;而属于青藏区的青海玛多县马胃蝇调查到2个种(黑腹胃蝇和鼻胃蝇);卡拉麦里与多伦县的马胃蝇组分相似性指数为1,二者与玛多县的相似性指数均为0.33; 3)多伦县的马胃蝇多样性指数(H'=1.31)显著大于玛多县(H'=0.23)以及新疆卡拉麦里(H'=0.21);宿主感染马胃蝇的均匀度指数(J)同样为多伦县(0.73)最高,其次为玛多县(0.34),卡拉麦里(0.12)最低; 4)黑腹胃蝇为卡拉麦里和玛多县马胃蝇的绝对优势种,分别占这2个地区采样总量的96.17%和93.17%;而在多伦县,肠胃蝇感染率最高(52.40%),其次为鼻胃蝇(23.80%),二者成为该地区的马胃蝇优势种。[结论] 1)蒙新区的马胃蝇种类较青藏区丰富; 2)多伦县具有丰富的马胃蝇类群; 3)马胃蝇多样性受宿主所在环境条件影响,蒙新区和青藏区马胃蝇物种的差异与2个地区巨大的自然条件差异有直接关系;干旱、荒漠以及高寒等恶劣生境下马胃蝇多样性低,其原因为某单一物种数量过大新疆或物种数少(青海); 4)黑腹胃蝇具有成功适应严酷生境的生存方式,黑腹胃蝇在干旱荒漠以及高寒环境感染量均占马胃蝇感染量的90%以上,与其与众不同的传播方式——将卵产于草上"污染"草场有着密切的联系。
关键词马胃蝇    感染率    多样性    蒙新区    青藏区    
Diversity and Infection of Gasterophilus spp. in Mongol-Xinjiang Region and Qinghai Tibet Region
Wang Wengting1, Xiao Sa1, Huang Heqing1, Li Kai1, Zhang Dong1, Chu Hongjun2, Guo Youqing3, Gao Wanli4    
1. College of Nature Conservation, Beijing Forestry University Beijing 100083;
2. Altay Management Station, Mt. Kalamaili Ungulate Nature Reserve Altay 836500;
3. Qingshan Forest Farm, Keshiketeng Banner, Chifeng, Inner Mongolia Chifeng 025366;
4. Forestry Department of Qinghai Xining 810008
Abstract: [Objective] A survey of infection in equids Gasterophilus spp. was conducted in Inner Mongol-Xinjiang region and Qinghai Tibet region in order to examine the infection diversity and parasitic characteristics of Gasterophilosis epidemic in the three regions.[Method] Equids in the Kalamaili Nature Reserve, Xinjiang, and Duolun, Inner Mongolia Autonomous Region in Mongol-Xinjiang region, Maduo, Qinghai Province in Qinghai Tibet region were treated with ivermectin to investigate and analyze the number and species of Gasterophilus spp. in the desert, desert-steppe and cold alpine steppe of these regions in the same winter.[Result] 1) Infection with Gasterophilus larvae was very common in horses in all these regions with an infection rate of 100%. 2) Gasterophilus spp. infecting the horses in both Xinjiang and Inner Mongolia belonged to six species of the genus, including G. pecorum, G. nasalis, G. intestinalis, G. nigricornis, G. haemorrhoidalis and G. inermis, which have been all reported in China, while only two species of the genus (G. pecorum and G. nasalis) were found in Qinghai. The component similarity index of Gasterophilus spp. was 1 (Xinjiang and Inner Mongolia), 0.33 (Xinjiang and Qinghai) and 0.33 (Inner Mongolia and Qinghai), respectively. 3) Shannon-Wiener index of Gasterophilus spp. that infected hosts was 1.31 in Duolun, Inner Mongolia, significantly higher than Maduo, Qinghai(0.23)and Kalamaili, Xinjiang (0.21), which also being part of Mongol-Xinjiang region. The evenness index of Gasterophilus spp. that infected hosts was highest in Duolun, Inner Mongolia (0.73), followed by that in Maduo, Qinghai (0.34), and lowest in Kalamaili, Xinjiang (0.12). This indicated a wider variety of Gasterophilus spp. species in Duolun, Inner Mongolia. 4) G. pecorum accounted for 96.17% and 93.17% of the larvae in Kalamaili, Xinjiang and Maduo, Qinghai, respectively, and was absolutely the dominant species in these two regions, while G. intestinalis (infection rate was 52.40%) and G. nasalis (infection rate was 23.80%) were the dominant species in Duolun, Inner Mongolia.[Conclusion] 1) The intensity of infection in Mongol-Xinjiang region was higher than that in Qinghai Tibet region. 2) More species were found in Mongol-Xinjiang region than Qinghai Tibet region. 3) The diversity of Gasterophilus spp. was influenced by the environmental conditions of its hosts. There would be a direct relationship between the differences of Gasterophilus species in Mongol-Xinjiang region and Qinghai Tibet region and the big differences of natural conditions of the two regions. Lower diversity was discovered under harsh conditions because there was oversize in some individual species or the small number of species. 4) G. pecorum would have successful reproductive strategies that adapts to the harsh environmental conditions. The prevalence of G. pecorum infection was more than 90% in both desert-steppe and cold alpine environment, and this might be closely related with its unique mode of transmission that laid eggs on grass and contaminated the pasture.
Key words: Gasterophilus spp.    infection rate    diversity    Mongol-Xinjiang region    Qinghai Tibet region    

马胃蝇属双翅目(Diptera)胃蝇科(Gasterophilidae)胃蝇属(Gasterophilus)昆虫,主要寄生于马科(Equidae)动物消化道内,1年1代,具有一定的专一性(Soulsby,1982)。幼虫在宿主消化道内发育至老熟幼虫,随宿主粪便排出,入地化蛹。马胃蝇在我国分布广泛,尤以东北、西北、内蒙古等草原地区以及云、贵、川等高海拨地区发生普遍(彭光政等,2011)。现今全球共记载9种马胃蝇(Zumpt,1965),其中黑腹胃蝇(G. pecorum)、肠胃蝇(G. intestinalis)、鼻胃蝇(G. nasalis)、黑角胃蝇(G. nigricornis)、红尾胃蝇(G. haemorrhoidalis)和裸节胃蝇(G. inermis)等6种我国已有记录(范滋德,1992王明福,1998)。

马胃蝇蛆病可使马科家畜役力、产肉、泌乳等生产性能下降,严重感染可致马匹衰竭死亡。在新疆卡拉麦里有蹄类自然保护区,马胃蝇蛆病流行于濒危物种、国家一级保护野生动物普氏野马(Equus przewalskii)群体中,且每年感染率高、程度重(张赫凡等,2008刘善辉,2012杨健梅等,2013王文婷等,2014),成为该物种种群恢复、健康发展的重要影响因子。卡拉麦里有6种马胃蝇,其优势种为黑腹胃蝇,蒙古野驴(E.hemionus)是普氏野马感染该虫的最主要疫源,该种马胃蝇因其宿主——蒙古野驴和牧民家马(E.caballus)生活空间的差异衍生出“荒漠型”和“山地型”2种遗传结构以及“山地型”分化于“荒漠型”(Wang et al., 2014)。

本研究以蒙新区、青藏区具有明显地域特点的新疆准格尔盆地卡拉麦里有蹄类自然保护区、内蒙古锡林郭勒盟多伦县和青海果洛藏族自治州玛多县的马科动物为研究对象,对上述3个地区的草原、荒漠草原和高寒草原马胃蝇蛆病发生情况及马胃蝇的多样性进行比较研究。

1 材料与方法 1.1 调查地点

卡拉麦里有蹄类自然保护区位于蒙新区西部荒漠亚区(88°30′—90°00′ E,44°38′—46°00′ N)面积170万hm2,海拔600~1 464 m。区内气候条件严酷,夏季炎热,冬季寒冷,年均气温2.4 ℃,年均降水量159 mm,年均蒸发量2 090 mm。保护区内地表水贫乏,主要水源为十几处泉水和低洼地积水(葛炎等,2003)。区域内马科动物包括常年生活于此地的蒙古野驴、普氏野马以及以此地为越冬场所的牧民家马(初红军等,2009)。

多伦县位于内蒙古锡林郭勒草原南端阴山北麓东端(115°30′—116°55′ E,41°45′—42°39′ N),海拔为1 150~1 800 m。锡林郭勒草原是世界驰名的四大草原之一,属欧亚大陆草原区,中温带半干旱向半湿润过渡的大陆性气候,年均气温1.6 ℃,年均降水量385 mm,年均蒸发量1 449.4~1 672.8 mm。多伦县草地面积16多万 hm2,占全县土地总面积的41.60%;当地马科动物为家马(阿如旱等,2007)。

玛多县位于青海省南部、果洛藏族自治州西北部(96°55′—99°20′ E,33°50′—35°40′ N),大部分地区海拔4 500~5 000 m,地形起伏不大,为高平原地区。气候属高原大陆性气候,年均气温-4.1 ℃,年均降水量303.9 mm,年均蒸发量为1 318. 86 mm(1961—1998);一年分为冬夏2季,冬季漫长而严寒,干燥多大风,夏季短促而温凉,多雨。玛多县草场面积220多万hm2,占全县总面积的87.5%;当地生活的马科动物主要是家马和藏野驴(E.kiang),其生活空间完全重叠(摆万奇等,2002)。

新疆、内蒙及青海是我国传统的牧区,在物种分布区划上分属于蒙新区和青藏区。本研究所选试验马匹均散放于当地草场,具有与当地马科动物广泛的接触机会,因此,卡拉麦里普氏野马马胃蝇感染情况与蒙古野驴和家马有关,玛多县试验家马感染情况与当地藏野驴和家马有联系,多伦县则受家马单一影响。

1.2 样品采集

分别于2014年2,3,4月在新疆卡拉麦里有蹄类自然保护区、内蒙多伦县、青海玛多县采集马胃蝇幼虫样品。王文婷等(2014)对黑腹胃蝇的研究显示,马胃蝇传播和感染与宿主种类没有关联性,因此,本研究以当地在上一个马胃蝇感染季以来未服药的散放马匹(野马、家马等)为采样对象。具体操作为: 在卡拉麦里随机圈取了9匹放归野马,在多伦县和玛多县随机各抽取了5匹春夏季自由游走于夏牧场的家马;给每匹马服用混有伊维菌素[北京万丰药业有限公司生产,兽药GMP证字(2006)278号]的饲料,每10 kg体质量投药剂量为1.0 g;连续4天收集粪便中排出的马胃蝇幼虫,并保存于无水乙醇中。幼虫样品经形态学分类鉴定(Zumpt,1965)后分别计数。

1.3 数据分析与处理

采用Shannon-Wiener多样性指数(H′)、Pielou均匀性指数(J)、Jaccard相似性指数(q)对3个地区马胃蝇多样性进行分析,相关计算公式如下:

$$H' = - \sum {{P_i}} \ln {P_i} \left( {i = 1,2,3, \cdots ,S} \right);$$ (1)
$$J = H'/H{'_{\max }};$$ (2)
$$q = N/\left( {{N_1} + {N_2} - N} \right).$$ (3)
式中为Pii种马胃蝇个体数与当地所有种马胃蝇个体总数的比值;S为物种数;H′max=ln SN为2个地区共有物种数,N1N2分别为地区1和地区2马胃蝇物种数(吴坤君等,2005; 张洁等,2010)。

2 结果与分析 2.1 不同地区宿主感染马胃蝇的数量及种类

3个地区共采集马胃蝇3龄幼虫12 257只,涉及6种马胃蝇(表 1),均为我国现有记录种。其中,新疆卡拉麦里和内蒙多伦县具有相同的马胃蝇种类,且遍及已知的6个种;青海玛多县采集的为黑腹胃蝇和鼻胃蝇2个种。对在4天采集时间段内收集到的马胃蝇数量进行分析发现,3个地区供试马匹马胃蝇的感染数量呈现出较大的倍数差异(表 1“单位采集量”),新疆卡拉麦里放归普氏野马的感染量显著高于内蒙古多伦县和青海玛多县,而内蒙古多伦县也明显高于青海玛多县的感染量。采样期间虽发生过一次短暂的鸦科鸟啄食粪便幼虫的情况,但幼虫损失数量按鸟啄食时间分析不足1/10。

表1 不同地区马胃蝇幼虫采集数量及种类 Tab. 1 Number and species of Gasterophilus larvae collected in different regions
2.2 不同地区宿主感染马胃蝇的组成

3个地区感染马胃蝇组成(表 2)显示,在新疆卡拉麦里和青海玛多县,黑腹胃蝇为绝对优势种,分别占各自地区采样总量的96.17%和93.17%,其他种类不足10%;在内蒙古多伦县,6种马胃蝇比例与前2个地区相比没有悬殊的差异,其主要感染物种也不同,肠胃蝇以52.40%居于首位,其次为鼻胃蝇(23.80%),黑腹胃蝇与红尾胃蝇相似,分别为9.42%和9.08%;黑角胃蝇和裸节胃蝇居后。

表2 不同地区宿主感染马胃蝇的组成 Tab. 2 Composition of Gasterophilus larvae collected in different regions
2.3 不同地区宿主感染马胃蝇的生物多样性指数

对3个地区宿主感染马胃蝇的多样性指数(H′)、均匀度指数(J)进行分析(表 3)可知,马胃蝇多样性指数(H′)排序为内蒙古多伦县(1.31)>青海玛多县(0.23)≈新疆卡拉麦里(0.21);马胃蝇均匀度指数(J)排序为内蒙古多伦县(0.73)>青海玛多县(0.34)>新疆卡拉麦里(0.12)。

表3 不同地区宿主感染马胃蝇的多样性 Tab. 3 Diversity of Gasterophilus larvae that infected hosts in different regions
2.4 不同地区宿主感染马胃蝇的组分相似性

3个地区宿主感染马胃蝇的组分相似性分析结果(表 4)表明,新疆卡拉麦里与内蒙多伦县的相似性指数为1,二者与青海玛多县的相似性指数均为0.33。

表4 不同地区宿主所感染马胃蝇组分相似性 Tab. 4 Similarity of composition of Gasterophilus larvae in different regions
3 结论与讨论

通过调查发现,3个地区马科动物均100%感染马胃蝇,类似情况在诸多研究中也均有报道(杨锡林,1986陈德明等,1987王善志等,1991李敬双等,1995彭光政等,2011),该现象显示。马胃蝇在进化过程中演化出独特而有效的运动能力、生存能力和寄生生存方式,从而根植于马科动物生存环境。

生物在与环境相互作用、协同进化的过程中,每个种都留下了深刻的环境烙印(Croizat,1952; Illies,1974; Brown,1988)。蒙新区和青藏区同属于古北界,受自然环境和条件的影响,形成了代表各自区系的特有动物种类,衍生出适应干旱荒漠草原生境的普氏野马和蒙古野驴以及适应高寒高原生境的藏野驴等马科动物类群(邓涛,1999; Connor et al., 2001; Turner,2004)。马胃蝇虽然专性寄生于马科动物,但其生活史中的蛹期和成虫阶段同样受环境因子影响而形成与本地环境及条件相应的生物种群。本研究中新疆卡拉麦里和内蒙多伦县马胃蝇物种数相同且种类一致表明,同属于蒙新区的二者间虽地跨多个地形,但其物种结构上仍符合一个物种板块的划定;在这方面,二者与青藏区玛多县仅有2个种的情况有明显差异。物种丰富度的地理格局主要是由能量对物种生理活动的直接控制引起的(Turner,2004),而良好的环境适合更多的物种生存(Currie,1991; Clarke et al., 2006)。因此,蒙新区和青藏区马胃蝇物种的差异与2个地区巨大的自然条件差异有直接关系。

优势种源自生存对策。在新疆卡拉麦里干旱荒漠草原以及青海玛多高寒草原,黑腹胃蝇高于90%的感染比例反映该物种演化出成功应对恶劣环境条件的生存对策,特点之一是其物种特殊的产卵机制——卵产于草上而非马体上(Zumpt,1965)。内蒙古多伦县毗邻广袤的锡林郭勒大草原,适宜的自然条件形成了该地区马胃蝇优势种为肠胃蝇和鼻胃蝇的结果,这类似于世界其他地区马胃蝇蛆病感染情况(Edwards,1982; Dunsmore et al., 1985; Principato,1989; Pandey et al., 1992; Reinemeyer et al., 2000; Sequeira et al., 2001; Felix et al., 2007; Studzińska et al., 2009)。

在马胃蝇物种丰富度方面,内蒙多伦县的多样性指数显著大于青海玛多县以及同属于蒙新区的新疆卡拉麦里,显示出内蒙多伦县具有丰富的马胃蝇类群及相对均衡的种群数量;新疆卡拉麦里马胃蝇群落尽管包含6个种,但其个体数量主要为黑腹胃蝇,以致多样性指数和均匀性指数较低,甚至低于青海玛多县。王文婷等(2014)对马胃蝇在马科动物宿主中并未形成遗传分化、没有宿主特异性的研究结论显示,蒙新区、青藏区马胃蝇分布格局的形成不受宿主种类的影响。因此,海拔、气候等环境条件对物种的分布及进化有着重要影响,该影响是通过环境热量实现的(Allen et al., 2007);青藏高原物种多样性很低与其高寒环境关系密切(方精云等,2004; 杨元合等,2004)。内蒙多伦县马胃蝇多样性指数及群落均匀度均高于同处于蒙新区的卡拉麦里,物种数更是高于青藏区的玛多县,与其适宜的降水量、温度等环境条件存在必然联系。

在马胃蝇感染量方面,与同处于蒙新区的多伦县相比,卡拉麦里普氏野马的感染量奇高,特别是黑腹胃蝇的感染数量。卡拉麦里气候与多伦县迥异,为干旱荒漠环境,加之黑腹胃蝇将卵产于草上,这与其他马胃蝇的传播方式不同,二者耦合成为该地区马胃蝇蛆病重度发生的首因。青海玛多县草原广袤,然而高寒的气候条件及短暂的夏季成为马胃蝇在宿主体外生存的限制条件,黑腹胃蝇虽然为绝对优势种,但马胃蝇感染总量与另外2地相比处于最低水平。

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