蝙蝠寄生革螨的分类研究
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摘要
蝙蝠体上的寄生革螨是我国尚待系统性研究的重要蝙蝠寄生虫类群,它们可能通过叮咬和寄主蝙蝠携带的微生物病原和病毒与包括人在内的高等动物发生直接或间接的重要关系,研究蝙蝠与寄生革螨间的协同进化关系,对认识动物-寄生物间协同进化关系以及物种演化机制具有重要意义。由于采集困难等客观因素的制约,蝙蝠寄生革螨系统学及其相关研究十分薄弱,有关中国区系的研究少而零散,亟待加强分类并拓展相关研究。为此,本文对中国蝙蝠寄生革螨区系进行了分类学研究,探讨了寄生革螨特定类群之属间和特定属之种间的系统发育关系、区系特征以及蝙蝠-寄生革螨间相互关系。
论文概述了蝙蝠体外寄生虫的构成类群及其分类地位,详细论述了革螨系统学研究历史、国内外研究情况、生物学特性,重点阐述了分类特征,比较了不同分类术语学。
研究材料采集自北京、天津、湖北、湖南、四川、重庆、贵州、广西、广东、福建及海南的200余个洞穴和约20座古建筑与民居栖息的蝙蝠。获得的宿主蝙蝠隶属于4科、11属、20种,寄生革螨标本从蝙蝠体表分离获得。
1.通过对蝙蝠寄生革螨的详细形态学比较研究和分类学研究,记述了中国蝙蝠寄生革螨2个科(巨刺螨科Macronyssidae Oudemans,1936;蝠螨科Spinturnicidae Oudemans,1902),6个属(浆刺螨属Ichoronyssus Kolenati,1858;巨刺螨属Macronyssus Kolenati,1858;肪刺螨属Steatonyssus Kolenati,1858;蝠螨属Spinturnix von Heyden,1826;拟弱螨属Paraperiglischrus Rudnick, 1960;埃螨属Eyndhovenia Rudnick,1960)共21种,其中包括5个新种,4个已知种雄螨首次描述,2个中国新记录种；绘制了准确反映新种、雄螨首次描述和已知种特征的形态图,编制了世界已知属和中国已知种的检索表。新种、首次雄螨记述种和中国新纪录种如下：
建立的5个新种隶属于巨刺螨科。隶属于巨刺螨属的4个新种分别为：拟雷氏巨刺螨Macronyssus pararadovskyi sp. nov.,以背板刚毛为27对,胸腺豆状、具粒状纹为主要区别特征；宿主是褐扁颅蝠Tylonycteris robustula (Thomas,1915)(广西)。武夷巨刺螨Macronyssus wuyiensis sp. nov.,以背板刚毛为21对,以位于生殖腹板边缘的Vz1及在板末端有1根副刚毛为主要鉴别特征；宿主是山蝠Nyctalus noctula Schreber,1774(福建)。山蝠巨刺螨Macronyssus nyctalus sp. Nov.,以雌螨背板具有22对刚毛、z4-z5长度比约为12.5：1,雄螨背板上具有J2为主要鉴别特征；宿主是山蝠Nyctalus noctula Schreber,1774(福建)。寡毛巨刺螨Macronyssus parachaetus sp. nov.,以体躯大,足细长,骨化弱,背板刚毛仅19对,j1位于背板的前缘表皮上,气门沟延伸到基节Ⅱ的中部,颚沟内具小齿12个为主要鉴别特征；宿主是大足鼠耳蝠Myotis ricketti (Thomas,1857)(北京)。隶属于肪刺螨属的1新种为吊罗肪刺螨Steatonyssus diaoluoensis sp. nov.,其以雌螨前背板具刚毛12对,后背板刚毛为10对；雄螨背板上具刚毛17对,腹面分为胸生殖腹板和肛板两部分,生殖毛gen.1根位于板外,1根位于板的边缘,其上还有4根腹毛为主要鉴别特征；宿主是小黄蝠Scotophilustemmincki(Horsfield,1824)(海南).
首次描述雄螨的4个种分别为肛拟弱螨Paraperiglischrus analis Pan et Teng,1973;来凤巨刺螨Macronyssus laifengensis Wang et Shi,1986;异棘巨刺螨Macronyssus miraspinosus Guet Wang,1985；(?)渡巨刺螨Macronyssus xianduensis(Zhou,Tang et Wen,1982)。
中国新记录2种分别为雷氏巨刺螨Macronyssus radovskyi (Domrow,1963);长翼蝠浆刺螨Ichoronyssus miniopteri Zumpt et Patterson,1952,,
2.本文应用PAUP*4.0系统发育软件,用NJ法和MP法对世界蝠螨科12个属和巨刺螨属48种进行了支序分析：
用34个特征构建蝠螨科12个属的系统树表明,颚湾蝠螨属Cameronieta和仿弱螨属Periglischrodes是姐妹群,这两个属和弱螨属Periglishchrus是姐妹群；但颚湾蝠螨属和弱螨属具有共同的祖征：气门沟很长,前端达到基节Ⅰ水平。钩螨属Oncosceles和距螨属Ancystropus是姐妹群,前面这5个属与拟弱螨属Paraperiglishchrus又是姐妹群,与本文作者主观推测的结果基本一致。蝠螨属Spinturnix和鞘尾蝠螨属Emballonuria是姐妹群；这2个属和拟蝠螨属Paraspinturnix又是姐妹群,与传统的观点相吻合,它们具有共同的衍征：背板骨化一般,全背板,气门沟较短,没有伸到腹部,仅延伸到基节Ⅱ和Ⅲ之间,胸叉退化,仅剩基部。拟裂螨属Parameristaspis形成一个单系,在蝠螨科的系统发育中处于基部的位置,是相对原始的属,其典型祖征有：背板骨化强,全背板,生殖腹板形状为舌形,胸板与生殖腹板贴近,胸板上的3对刚毛都位于胸板上。
用38个特征对巨刺螨属48种进行支序分析表明,两种方法的结果差别较大,MP法能大致把本属48个种分为5个种团(species group),而NJ法大致分为6个种团,除了jonesi, cavus, japonicus, parachaetus, ellipticus, longimanus, cyclaspis这个种团较稳定外,它们具有共同的衍征：体长大于800μm,躯体长大于颚体长的3倍,颚体长度大于或等于足Ⅱ位置躯体的宽度,背板小,仅覆盖背面大于等于1/3,小于1/2,板上的刚毛数小于等于23对,气门沟前端延伸到基节Ⅱ。其它种团都有一些变化,所以以目前的分析结果表明,本属不适合分为种团,可能原因有多种：特征的极性,数量,各个种间的特征交叉的太多等。
3.对蝠螨科各属的区系特征分析表明,世界现知12属中以东洋区属最为丰富,新北区、东洋区和新热带区各自有本区的特有属,说明蝠螨科虽是世界性分布的类群,但其属级单元的地理分布具有明显地理区域性。对巨刺螨科巨刺螨属现知48种和4个新种的区系特征分析表明,其以东洋区种和古北区种最为丰富,而世界六大地理区都有该属的特有种,说明巨刺螨属虽是世界性分布的类群,但该属种级单元的地理分布具有明显地理区域性。对肪刺螨属现知49种和1个新种的区系特征分析表明,其古北区、东洋区和非洲区的种类最为丰富,六大地理区都有其特有种,说明肪刺螨属虽是世界性分布的类群,但其种级单元的地理分布具有明显地理区域性。但这些突出区系特点可能和各地理区系研究尚欠深度和广度有关。
4.分析讨论了20种蝙蝠和21种寄生革螨间的相互关系,本文认为寄主蝙蝠与寄生革螨间具有相对稳定的寄生专一性关系,并将之概括为革螨与其寄主蝙蝠间的“属-属”对应关系和“种-种”对应关系。特定的寄生革螨类群具有各自的寄生生物学特性,其功能形态特征的进化与之相适应。
The ectoparasitic gamasid mites of bats are distributed widely with the hosts, and there are 2 families,8 genera,46 previously described species from China up to now. It is the first preliminary study covering morphology, taxonomy, cladistics on the ectoparasitic gamasid mites of bats from China.
The classification history, taxonomic status and general bionomics of the gamasid mite are reviewed, and morphological characters and terminology used for taxonomic study of the group are discussed in detail in the present paper.
The bats are collected from more than 200 caves and about 20 ancient and residential buildings in Beijing, Tianjin, Hubei, Hunan, Sichuan, Chongqing, Guizhou, Guangxi, Guangdong, Fujian and Hainan. The host bats belong to 4 families,11 genera and 20 species, from which parasitic gamasid mite specimens are isolated.
1. There are 21 species of the ectoparasitic gamasid mites of bats identified and described. They belong to two families (Macronyssidae Oudemans,1936 and Spinturnicidae Oudemans, 1902), five genera (Macronyssus Kolenati,1858; Steatonyssus Kolenati,1858; Ichoronyssus Kolenati,1858; Spinturnix von Heyden,1826; Paraperiglischrus Rudnick,1960; Eyndhovenia Rudnick,1960), of which 2 are newly recorded species from China,4 known species first described with males,5 species are newly to science. They are:M. pararadovskyi sp. nov., M. wuyiensis sp. nov., Macronyssus nyctalus sp. nov., M. parachaetus sp. nov., Steatonyssus diaoluoensis sp. nov.. The males of Paraperiglischrus analis Pan et Teng,1973, M. laifengensis Wang et Shi,1986, Macronyssus miraspinosus Gu et Wang,1985, Macronyssus xianduensis (Zhou, Tang et Wen,1982) are described for first time.2 species firstly record from China are Macronyssus radovskyi (Domrow,1963) and Ichoronyssus miniopteri Zumpt et Patterson,1952. It states diagnosis characters of 2 families,5 genera,21 known species primary diagnosis features, with described and illustrated.
The 5 new species belong to the Family Macronyssidae Oudemans,1936 and their main distinguishable characters from the close species and the hosts are as follows:In M. pararadovskyi sp. nov., dorsal plate with 27 pairs of setae, sternal glands fabaceous, bearing granulated stripe; the host is Tylonycteris robustula (Thomas,1915) (Guangxi). In M. wuyiensis sp. nov., dorsal plate with 21 pairs of setae, Zvl lying at the margin of the epigynial plate and the posterior with 1 other seta; the host is Nyctalus noctula Schreber,1774 (Fujian). In Macronyssus nyctalus sp. nov., female dorsal plate with 22 pairs of setae, z4-z5 length ratio 12.5:1, male dorsal plate with J2 present; and the host is N. noctula Schreber,1774 (Fujian). In M. parachaetus sp. nov., idiosoma large, weakly sclerotized, legs slender and long, dorsal plate with only 19 pairs of setae, j1 anterior to the plate, peritreme terminating over posterior one-half or about the middle of coxa II, plate extending anteriorly as narrow strip which widens and then narrows to termination over coxa I, deutosternal groove with 12 denticles arranged 1-1-2-1-1-1-2-1-1-1; the host is Myotis ricketti (Thomas,1857) (Beijing). In Steatonyssus diaoluoensis sp. nov., female podosomal plate with 12 pairs of setae, opisthosomal plate bearing 10 pairs of primary setae; male dorsal plate with 17 pairs of setae, ventral armature divided into sterno-genito-ventral plate and anal plate, the foreplate bearing st.1-st.3, pair of gen.(one on unarmed integument, one on margin of plate) and four other setae; the host is Scotophilus temmincki (Horsfield,1824) (Hainan).
2. Cladistic analysis of 12 genera of Spinturnicidae and 48 species of Macronyssus based on female specimens and with the genus Ichoronyssus and Ichoronyssus scutatus as the outgroups were performed using method NJ and MP of PAUP*4.0 program.
The phylogenetic trees for 12 genera of Spinturnicidae built using 34 characteristics show that Cameronieta and Periglischrodes are sister group with same ancestral characters:peritreme long and terminating over coxa I; moreover the two genera and Periglishchrus are sister group, Oncosceles and Ancystropus are sister group, too, all the former 5 genera are sister group with Paraperiglishchrus, it is according to our expectation. Spinturnix and Emballonuria is sister group, the two genera are sister group with Paraspinturnix, this is inosculating as traditionary viewpoint, with same derived characters:dorsal plate entirely, moderate sclerotized, peritreme short, completely dorsal, and terminating over coxaⅡ-Ⅲ, tritosternum reduced, subcircular, lightly sclerotized. Parameristaspis is a monophyletic, it is placed in the base of Spinturnicidae, the ancestral characters:dorsal plate entirely, strong sclerotized, st.1-st.3 on the sternal plate, genital plate tongue-like.
The phylogenetic trees for 48 species of the genus Macronysssus built using 38 characteristics show that:with MP, it divided the Macronysssus into 5 species groups, but with NJ, it divided 6 species groups. Only jonesi, cavus, japonicus, parachaetus, ellipticus, longimanus, cyclaspis groups steadily, they have same derived characters:idiosoma large, over 800μm, length of idiosoma and length of gnathosoma ratio more over 3:1, length of idiosoma no less than the width of idiosoma at the leg II, dorsal plate small, only cover with about 1/3 dorsum, with no more than 23 pairs of setae, peritreme terminating over coxa II. So it suggested that species groups could be not applicable for sub-classification of Macronyssus until more materials and information available.
3. The analysis of faunal characteristics suggest that Oriental taxa make up the bulk of 12 genera known worldwide, and the Neoarctic, Neotropical and Oriental regions have their endemic genera respectively. This indicates that the family is widely distributed one, the distributions of the taxa at generic level owns regional characteristics. The faunal analysis on 48 species and 4 new species of Macronyssus show that members originated of Oriental and Palaearctic region abundant, and all six biogeographic regions have the endemic species. This means that although it is a genus distributed world-wide group, the taxa at species level have regional features. The faunal analysis on 49 species and 1 new species of Steatonyssus show that members originated of Oriental, Palaearctic region and Ethiopian region abundant, and all six biogeographic regions have the endemic species. This means that although it is a genus distributed world-wide group, the taxa at species level have regional features.
4. Host-specificity among the 21 gamasid mite species and 20 bat species are discussed the analyzed. It could be inferred that there is a relatively stable specific relationship among the host bats and the parasitic mites. The relationships of host-mite are proposed as‘genus-genus’pattern and'species-species’pattern in the present paper. The every gamasid group has developed its own parasitic-biological characteristics, and correspondingly has specific morphologic characteristics representing adaptive evolution of the functional structure.
论文概述了蝙蝠体外寄生虫的构成类群及其分类地位,详细论述了革螨系统学研究历史、国内外研究情况、生物学特性,重点阐述了分类特征,比较了不同分类术语学。
研究材料采集自北京、天津、湖北、湖南、四川、重庆、贵州、广西、广东、福建及海南的200余个洞穴和约20座古建筑与民居栖息的蝙蝠。获得的宿主蝙蝠隶属于4科、11属、20种,寄生革螨标本从蝙蝠体表分离获得。
1.通过对蝙蝠寄生革螨的详细形态学比较研究和分类学研究,记述了中国蝙蝠寄生革螨2个科(巨刺螨科Macronyssidae Oudemans,1936;蝠螨科Spinturnicidae Oudemans,1902),6个属(浆刺螨属Ichoronyssus Kolenati,1858;巨刺螨属Macronyssus Kolenati,1858;肪刺螨属Steatonyssus Kolenati,1858;蝠螨属Spinturnix von Heyden,1826;拟弱螨属Paraperiglischrus Rudnick, 1960;埃螨属Eyndhovenia Rudnick,1960)共21种,其中包括5个新种,4个已知种雄螨首次描述,2个中国新记录种；绘制了准确反映新种、雄螨首次描述和已知种特征的形态图,编制了世界已知属和中国已知种的检索表。新种、首次雄螨记述种和中国新纪录种如下：
建立的5个新种隶属于巨刺螨科。隶属于巨刺螨属的4个新种分别为：拟雷氏巨刺螨Macronyssus pararadovskyi sp. nov.,以背板刚毛为27对,胸腺豆状、具粒状纹为主要区别特征；宿主是褐扁颅蝠Tylonycteris robustula (Thomas,1915)(广西)。武夷巨刺螨Macronyssus wuyiensis sp. nov.,以背板刚毛为21对,以位于生殖腹板边缘的Vz1及在板末端有1根副刚毛为主要鉴别特征；宿主是山蝠Nyctalus noctula Schreber,1774(福建)。山蝠巨刺螨Macronyssus nyctalus sp. Nov.,以雌螨背板具有22对刚毛、z4-z5长度比约为12.5：1,雄螨背板上具有J2为主要鉴别特征；宿主是山蝠Nyctalus noctula Schreber,1774(福建)。寡毛巨刺螨Macronyssus parachaetus sp. nov.,以体躯大,足细长,骨化弱,背板刚毛仅19对,j1位于背板的前缘表皮上,气门沟延伸到基节Ⅱ的中部,颚沟内具小齿12个为主要鉴别特征；宿主是大足鼠耳蝠Myotis ricketti (Thomas,1857)(北京)。隶属于肪刺螨属的1新种为吊罗肪刺螨Steatonyssus diaoluoensis sp. nov.,其以雌螨前背板具刚毛12对,后背板刚毛为10对；雄螨背板上具刚毛17对,腹面分为胸生殖腹板和肛板两部分,生殖毛gen.1根位于板外,1根位于板的边缘,其上还有4根腹毛为主要鉴别特征；宿主是小黄蝠Scotophilustemmincki(Horsfield,1824)(海南).
首次描述雄螨的4个种分别为肛拟弱螨Paraperiglischrus analis Pan et Teng,1973;来凤巨刺螨Macronyssus laifengensis Wang et Shi,1986;异棘巨刺螨Macronyssus miraspinosus Guet Wang,1985；(?)渡巨刺螨Macronyssus xianduensis(Zhou,Tang et Wen,1982)。
中国新记录2种分别为雷氏巨刺螨Macronyssus radovskyi (Domrow,1963);长翼蝠浆刺螨Ichoronyssus miniopteri Zumpt et Patterson,1952,,
2.本文应用PAUP*4.0系统发育软件,用NJ法和MP法对世界蝠螨科12个属和巨刺螨属48种进行了支序分析：
用34个特征构建蝠螨科12个属的系统树表明,颚湾蝠螨属Cameronieta和仿弱螨属Periglischrodes是姐妹群,这两个属和弱螨属Periglishchrus是姐妹群；但颚湾蝠螨属和弱螨属具有共同的祖征：气门沟很长,前端达到基节Ⅰ水平。钩螨属Oncosceles和距螨属Ancystropus是姐妹群,前面这5个属与拟弱螨属Paraperiglishchrus又是姐妹群,与本文作者主观推测的结果基本一致。蝠螨属Spinturnix和鞘尾蝠螨属Emballonuria是姐妹群；这2个属和拟蝠螨属Paraspinturnix又是姐妹群,与传统的观点相吻合,它们具有共同的衍征：背板骨化一般,全背板,气门沟较短,没有伸到腹部,仅延伸到基节Ⅱ和Ⅲ之间,胸叉退化,仅剩基部。拟裂螨属Parameristaspis形成一个单系,在蝠螨科的系统发育中处于基部的位置,是相对原始的属,其典型祖征有：背板骨化强,全背板,生殖腹板形状为舌形,胸板与生殖腹板贴近,胸板上的3对刚毛都位于胸板上。
用38个特征对巨刺螨属48种进行支序分析表明,两种方法的结果差别较大,MP法能大致把本属48个种分为5个种团(species group),而NJ法大致分为6个种团,除了jonesi, cavus, japonicus, parachaetus, ellipticus, longimanus, cyclaspis这个种团较稳定外,它们具有共同的衍征：体长大于800μm,躯体长大于颚体长的3倍,颚体长度大于或等于足Ⅱ位置躯体的宽度,背板小,仅覆盖背面大于等于1/3,小于1/2,板上的刚毛数小于等于23对,气门沟前端延伸到基节Ⅱ。其它种团都有一些变化,所以以目前的分析结果表明,本属不适合分为种团,可能原因有多种：特征的极性,数量,各个种间的特征交叉的太多等。
3.对蝠螨科各属的区系特征分析表明,世界现知12属中以东洋区属最为丰富,新北区、东洋区和新热带区各自有本区的特有属,说明蝠螨科虽是世界性分布的类群,但其属级单元的地理分布具有明显地理区域性。对巨刺螨科巨刺螨属现知48种和4个新种的区系特征分析表明,其以东洋区种和古北区种最为丰富,而世界六大地理区都有该属的特有种,说明巨刺螨属虽是世界性分布的类群,但该属种级单元的地理分布具有明显地理区域性。对肪刺螨属现知49种和1个新种的区系特征分析表明,其古北区、东洋区和非洲区的种类最为丰富,六大地理区都有其特有种,说明肪刺螨属虽是世界性分布的类群,但其种级单元的地理分布具有明显地理区域性。但这些突出区系特点可能和各地理区系研究尚欠深度和广度有关。
4.分析讨论了20种蝙蝠和21种寄生革螨间的相互关系,本文认为寄主蝙蝠与寄生革螨间具有相对稳定的寄生专一性关系,并将之概括为革螨与其寄主蝙蝠间的“属-属”对应关系和“种-种”对应关系。特定的寄生革螨类群具有各自的寄生生物学特性,其功能形态特征的进化与之相适应。
The ectoparasitic gamasid mites of bats are distributed widely with the hosts, and there are 2 families,8 genera,46 previously described species from China up to now. It is the first preliminary study covering morphology, taxonomy, cladistics on the ectoparasitic gamasid mites of bats from China.
The classification history, taxonomic status and general bionomics of the gamasid mite are reviewed, and morphological characters and terminology used for taxonomic study of the group are discussed in detail in the present paper.
The bats are collected from more than 200 caves and about 20 ancient and residential buildings in Beijing, Tianjin, Hubei, Hunan, Sichuan, Chongqing, Guizhou, Guangxi, Guangdong, Fujian and Hainan. The host bats belong to 4 families,11 genera and 20 species, from which parasitic gamasid mite specimens are isolated.
1. There are 21 species of the ectoparasitic gamasid mites of bats identified and described. They belong to two families (Macronyssidae Oudemans,1936 and Spinturnicidae Oudemans, 1902), five genera (Macronyssus Kolenati,1858; Steatonyssus Kolenati,1858; Ichoronyssus Kolenati,1858; Spinturnix von Heyden,1826; Paraperiglischrus Rudnick,1960; Eyndhovenia Rudnick,1960), of which 2 are newly recorded species from China,4 known species first described with males,5 species are newly to science. They are:M. pararadovskyi sp. nov., M. wuyiensis sp. nov., Macronyssus nyctalus sp. nov., M. parachaetus sp. nov., Steatonyssus diaoluoensis sp. nov.. The males of Paraperiglischrus analis Pan et Teng,1973, M. laifengensis Wang et Shi,1986, Macronyssus miraspinosus Gu et Wang,1985, Macronyssus xianduensis (Zhou, Tang et Wen,1982) are described for first time.2 species firstly record from China are Macronyssus radovskyi (Domrow,1963) and Ichoronyssus miniopteri Zumpt et Patterson,1952. It states diagnosis characters of 2 families,5 genera,21 known species primary diagnosis features, with described and illustrated.
The 5 new species belong to the Family Macronyssidae Oudemans,1936 and their main distinguishable characters from the close species and the hosts are as follows:In M. pararadovskyi sp. nov., dorsal plate with 27 pairs of setae, sternal glands fabaceous, bearing granulated stripe; the host is Tylonycteris robustula (Thomas,1915) (Guangxi). In M. wuyiensis sp. nov., dorsal plate with 21 pairs of setae, Zvl lying at the margin of the epigynial plate and the posterior with 1 other seta; the host is Nyctalus noctula Schreber,1774 (Fujian). In Macronyssus nyctalus sp. nov., female dorsal plate with 22 pairs of setae, z4-z5 length ratio 12.5:1, male dorsal plate with J2 present; and the host is N. noctula Schreber,1774 (Fujian). In M. parachaetus sp. nov., idiosoma large, weakly sclerotized, legs slender and long, dorsal plate with only 19 pairs of setae, j1 anterior to the plate, peritreme terminating over posterior one-half or about the middle of coxa II, plate extending anteriorly as narrow strip which widens and then narrows to termination over coxa I, deutosternal groove with 12 denticles arranged 1-1-2-1-1-1-2-1-1-1; the host is Myotis ricketti (Thomas,1857) (Beijing). In Steatonyssus diaoluoensis sp. nov., female podosomal plate with 12 pairs of setae, opisthosomal plate bearing 10 pairs of primary setae; male dorsal plate with 17 pairs of setae, ventral armature divided into sterno-genito-ventral plate and anal plate, the foreplate bearing st.1-st.3, pair of gen.(one on unarmed integument, one on margin of plate) and four other setae; the host is Scotophilus temmincki (Horsfield,1824) (Hainan).
2. Cladistic analysis of 12 genera of Spinturnicidae and 48 species of Macronyssus based on female specimens and with the genus Ichoronyssus and Ichoronyssus scutatus as the outgroups were performed using method NJ and MP of PAUP*4.0 program.
The phylogenetic trees for 12 genera of Spinturnicidae built using 34 characteristics show that Cameronieta and Periglischrodes are sister group with same ancestral characters:peritreme long and terminating over coxa I; moreover the two genera and Periglishchrus are sister group, Oncosceles and Ancystropus are sister group, too, all the former 5 genera are sister group with Paraperiglishchrus, it is according to our expectation. Spinturnix and Emballonuria is sister group, the two genera are sister group with Paraspinturnix, this is inosculating as traditionary viewpoint, with same derived characters:dorsal plate entirely, moderate sclerotized, peritreme short, completely dorsal, and terminating over coxaⅡ-Ⅲ, tritosternum reduced, subcircular, lightly sclerotized. Parameristaspis is a monophyletic, it is placed in the base of Spinturnicidae, the ancestral characters:dorsal plate entirely, strong sclerotized, st.1-st.3 on the sternal plate, genital plate tongue-like.
The phylogenetic trees for 48 species of the genus Macronysssus built using 38 characteristics show that:with MP, it divided the Macronysssus into 5 species groups, but with NJ, it divided 6 species groups. Only jonesi, cavus, japonicus, parachaetus, ellipticus, longimanus, cyclaspis groups steadily, they have same derived characters:idiosoma large, over 800μm, length of idiosoma and length of gnathosoma ratio more over 3:1, length of idiosoma no less than the width of idiosoma at the leg II, dorsal plate small, only cover with about 1/3 dorsum, with no more than 23 pairs of setae, peritreme terminating over coxa II. So it suggested that species groups could be not applicable for sub-classification of Macronyssus until more materials and information available.
3. The analysis of faunal characteristics suggest that Oriental taxa make up the bulk of 12 genera known worldwide, and the Neoarctic, Neotropical and Oriental regions have their endemic genera respectively. This indicates that the family is widely distributed one, the distributions of the taxa at generic level owns regional characteristics. The faunal analysis on 48 species and 4 new species of Macronyssus show that members originated of Oriental and Palaearctic region abundant, and all six biogeographic regions have the endemic species. This means that although it is a genus distributed world-wide group, the taxa at species level have regional features. The faunal analysis on 49 species and 1 new species of Steatonyssus show that members originated of Oriental, Palaearctic region and Ethiopian region abundant, and all six biogeographic regions have the endemic species. This means that although it is a genus distributed world-wide group, the taxa at species level have regional features.
4. Host-specificity among the 21 gamasid mite species and 20 bat species are discussed the analyzed. It could be inferred that there is a relatively stable specific relationship among the host bats and the parasitic mites. The relationships of host-mite are proposed as‘genus-genus’pattern and'species-species’pattern in the present paper. The every gamasid group has developed its own parasitic-biological characteristics, and correspondingly has specific morphologic characteristics representing adaptive evolution of the functional structure.
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