刺触螨科系统学及其形态与分子系统发育的研究
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摘要
刺触螨科隶属于蜱螨亚纲Acari，辐螨总目Superorder Actinotrichida，前气门目Prostigmata，寄殖螨亚目Suborder Parasitengona，水螨群Phalanx Hydracnidia，腺水螨总科Lebertioidea，是区系研究薄弱，分类地位存在分歧的类群。本文对中国刺触螨科区系进行了分类学研究，并从形态学和分子生物学两方面分析了刺触螨及其近缘类群间的系统发育关系。
本文详细总结论述了蜱螨学与水螨系统学的发展、刺触螨科的分类地位以及分类学研究历史；介绍了分子系统学的概念、分子系统学常用的研究方法及其在螨类系统学研究中的应用。
经形态和分类研究，发现并记述了1个中国新记录属(拟刺触螨属)、1个中国新记录亚属(短须剌触螨亚属)、3个中国新纪录种和1个新种；共记述刺触螨科1亚科(刺触螨亚科)2属(刺触螨属、拟刺触螨属)3亚属(鬃刺触螨亚属、刺触螨亚属、短须刺触螨亚属)15种；编制了刺触螨科属(亚属)和中国已知属、种的分类检索表。
选用22个形态学特征，应用PAUP~*4.0的MP和NJ分析方法，探讨了刺触螨科内属(亚属)间系统发育关系，刺触螨和腺水螨总科下另外7科的系统发育关系。刺触螨科内属(亚属)间系统发育关系结果显示与Cook(1974)的主观推导相似。但是原来划分为刺触螨亚科下的伊利刺触螨亚属Illiesiella与安迪刺触螨属Andinosperchon的关系似乎更加密切，在树中聚为一支，似乎应作为脊刺触螨属Notosperchonopsis一个亚属单元。原来划归刺触螨属Sperchon的阿卡刺触螨亚属Acadiosperchon与脊刺触螨亚属Notosperchonopsis关系更为密切，而似乎也应该将之作为脊刺触螨厲Notosperchonopsis一个亚属单元。本文据此提出了刺触螨2亚科、6属及其中2属各含4个亚属的系统分类建议。
选用23个形态学特征和2个生物学特征，对刺触螨和腺水螨总科下另外7科的系统发育关系分析结果基本和Cook(1974)分类系统相一致，但构建的系统树显示，刺触螨应具有总科的分类地位，即将刺触螨科从腺水螨总科中划分出来，建立独立的刺触螨总科Sperchonioidea。
详细探索了水螨基因组提取、PCR扩增、转化到测序的系列方法，经技术改进，成功扩增出阔毛曲跗湿螨的COⅠ基因片段、寡毛刺触螨和三叉腺水螨的28S rDNA D3基因片段；其中28S rDNA D3为首次从水螨群中获得。用PAUP~*4.0的MP、ML、NJ法，对所得的阔毛曲跗湿螨的COⅠ基因片段和从GeneBank下载的7条同源序列进行系统发育分析，结果和Cook(1974)分类系统相近，不支持将刺触螨科提升为总科，但由于水螨注册序列资料有限，这一结果尚待进一步研究检验。
Sperchontidae is a group, less known from China, of Lebertioidea, Phalanx Hydracnidia, Suborder Parasitengona, Prostigmata, Superorder Actinotrichida, Acari.. Some acarologists suggested that Sperchontidae should be classified as a single superfamily Sperchonioidea. The present works covered taxonomics of the family from China and phylogenetic relationships within the group and among the families of Lebertioidea and related others based on both the morphological and molecular approaches. The paper includes 4 chapters:
Chapter One briefly reviewed the development of Acarology, studies in Hydrachnidia, and the classification history of Sperchontidae. Advance in molecular phylogeny of mite are introduced with potential approach.
In Chapter Two, 15 species of the Sperchontidae are described. They belong to the subfamily Sperchoninae, two genera (Sperchonopsis and Sperchon), three subgenus {Palpisperchon, Hispidosperchon and Sperchon) separately, of them 1 species is new to science, Sperchon(Hispidosperchon)jianfengensis sp. nov., and 3 newly recorded from China, Sperchonopsis echphyma, Sperchon (Palpisperchon) mirabilis and Sperchon (Hispidosperchon) placoderma. Keys to the genera and to the known species from China are given.
Chapter Three focused on reconstructing of phylogenetic relationships, by means of cladistic analysis using 22 morphological characters, among genera and subgenera in Sperchontidae and families in Lebertioidea and close superfmilies. The result showed that phylogenetic relationships of 11 Sperchontidae genera and subgenera do not agree with that proposed by Cook (1974). Acadiosperchon should be treated as a subgenus of Notosperchonopsis; genera Illiesiella an Andinosperchon were sister groups and should be treated as two subgenera of Notosperchonopsis. Relationships of with the others consist with that proposed by Cook (1980). Hence, the present paper proposed a phylogenetic scheme with 2 subfamilies, 6 genera, of which 2 genera each contained 4 subgenera.
The phylogenetic relationships of 8 Lebertioidea families, based on the cladistic analysis made using 23 morphological and 2 biology characters, suggested by Cook (1974) seems not agree with the nature. And the cladgram showed that Sperchontidae seemed not a family of Lebertioidea, but an independent superfamily from Lebertioidea.
The methods for extracting of DNA were compared for optimizing conditions of CTAB, Chelex-100 and PCR. By a modification method of the CTAB. Fragments of mtDNA CO I gene in Atractides (Atractides) latisetus and fragments of 28S rDNA D3 gene in Sperchon (Hispidosperchon) oligospinis and Lebertia (Pilolebeitia) trifurcilla were sequenced. The homologous sequences for mtDNA CO I of 7 species of mites were downloaded from the GenBank data library. Phylogenetic tree were reconstructed using the sequences of mtDNA COI of the eight species with MP, ML and NJ of software PAUP*4.0. The result showed that Sperchontidae should not be an independent superfamily in hierarchy, but further examining of the conclusion should be made once there are more available sequences of the fragment.
本文详细总结论述了蜱螨学与水螨系统学的发展、刺触螨科的分类地位以及分类学研究历史；介绍了分子系统学的概念、分子系统学常用的研究方法及其在螨类系统学研究中的应用。
经形态和分类研究，发现并记述了1个中国新记录属(拟刺触螨属)、1个中国新记录亚属(短须剌触螨亚属)、3个中国新纪录种和1个新种；共记述刺触螨科1亚科(刺触螨亚科)2属(刺触螨属、拟刺触螨属)3亚属(鬃刺触螨亚属、刺触螨亚属、短须刺触螨亚属)15种；编制了刺触螨科属(亚属)和中国已知属、种的分类检索表。
选用22个形态学特征，应用PAUP~*4.0的MP和NJ分析方法，探讨了刺触螨科内属(亚属)间系统发育关系，刺触螨和腺水螨总科下另外7科的系统发育关系。刺触螨科内属(亚属)间系统发育关系结果显示与Cook(1974)的主观推导相似。但是原来划分为刺触螨亚科下的伊利刺触螨亚属Illiesiella与安迪刺触螨属Andinosperchon的关系似乎更加密切，在树中聚为一支，似乎应作为脊刺触螨属Notosperchonopsis一个亚属单元。原来划归刺触螨属Sperchon的阿卡刺触螨亚属Acadiosperchon与脊刺触螨亚属Notosperchonopsis关系更为密切，而似乎也应该将之作为脊刺触螨厲Notosperchonopsis一个亚属单元。本文据此提出了刺触螨2亚科、6属及其中2属各含4个亚属的系统分类建议。
选用23个形态学特征和2个生物学特征，对刺触螨和腺水螨总科下另外7科的系统发育关系分析结果基本和Cook(1974)分类系统相一致，但构建的系统树显示，刺触螨应具有总科的分类地位，即将刺触螨科从腺水螨总科中划分出来，建立独立的刺触螨总科Sperchonioidea。
详细探索了水螨基因组提取、PCR扩增、转化到测序的系列方法，经技术改进，成功扩增出阔毛曲跗湿螨的COⅠ基因片段、寡毛刺触螨和三叉腺水螨的28S rDNA D3基因片段；其中28S rDNA D3为首次从水螨群中获得。用PAUP~*4.0的MP、ML、NJ法，对所得的阔毛曲跗湿螨的COⅠ基因片段和从GeneBank下载的7条同源序列进行系统发育分析，结果和Cook(1974)分类系统相近，不支持将刺触螨科提升为总科，但由于水螨注册序列资料有限，这一结果尚待进一步研究检验。
Sperchontidae is a group, less known from China, of Lebertioidea, Phalanx Hydracnidia, Suborder Parasitengona, Prostigmata, Superorder Actinotrichida, Acari.. Some acarologists suggested that Sperchontidae should be classified as a single superfamily Sperchonioidea. The present works covered taxonomics of the family from China and phylogenetic relationships within the group and among the families of Lebertioidea and related others based on both the morphological and molecular approaches. The paper includes 4 chapters:
Chapter One briefly reviewed the development of Acarology, studies in Hydrachnidia, and the classification history of Sperchontidae. Advance in molecular phylogeny of mite are introduced with potential approach.
In Chapter Two, 15 species of the Sperchontidae are described. They belong to the subfamily Sperchoninae, two genera (Sperchonopsis and Sperchon), three subgenus {Palpisperchon, Hispidosperchon and Sperchon) separately, of them 1 species is new to science, Sperchon(Hispidosperchon)jianfengensis sp. nov., and 3 newly recorded from China, Sperchonopsis echphyma, Sperchon (Palpisperchon) mirabilis and Sperchon (Hispidosperchon) placoderma. Keys to the genera and to the known species from China are given.
Chapter Three focused on reconstructing of phylogenetic relationships, by means of cladistic analysis using 22 morphological characters, among genera and subgenera in Sperchontidae and families in Lebertioidea and close superfmilies. The result showed that phylogenetic relationships of 11 Sperchontidae genera and subgenera do not agree with that proposed by Cook (1974). Acadiosperchon should be treated as a subgenus of Notosperchonopsis; genera Illiesiella an Andinosperchon were sister groups and should be treated as two subgenera of Notosperchonopsis. Relationships of with the others consist with that proposed by Cook (1980). Hence, the present paper proposed a phylogenetic scheme with 2 subfamilies, 6 genera, of which 2 genera each contained 4 subgenera.
The phylogenetic relationships of 8 Lebertioidea families, based on the cladistic analysis made using 23 morphological and 2 biology characters, suggested by Cook (1974) seems not agree with the nature. And the cladgram showed that Sperchontidae seemed not a family of Lebertioidea, but an independent superfamily from Lebertioidea.
The methods for extracting of DNA were compared for optimizing conditions of CTAB, Chelex-100 and PCR. By a modification method of the CTAB. Fragments of mtDNA CO I gene in Atractides (Atractides) latisetus and fragments of 28S rDNA D3 gene in Sperchon (Hispidosperchon) oligospinis and Lebertia (Pilolebeitia) trifurcilla were sequenced. The homologous sequences for mtDNA CO I of 7 species of mites were downloaded from the GenBank data library. Phylogenetic tree were reconstructed using the sequences of mtDNA COI of the eight species with MP, ML and NJ of software PAUP*4.0. The result showed that Sperchontidae should not be an independent superfamily in hierarchy, but further examining of the conclusion should be made once there are more available sequences of the fragment.
引文
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64. Edwardsa, O.R, Melob, E.L, Smith, L, M.A. Hoy. Discrimination of three Typhlodromalus species(Acari: Phytoseiidae) using random amplified polymorphic DNA markers. Exp Appl Acarol,1997, 21: 101-109.
65. Ernsting, B.R, Edwards, D.D, Vidrine, M.F, Myers,K.S, Harmon,C.M. Phylogenetic relationships among species of the subgenus Parasitatax(Acari: Unionicolidae: Unionicola) based on DNA sequence of the mitochondrial cytochrome oxidase I gene. International Journal Of Acarology,2006,32(2): 195-202.
66. Essig, A, Rinder, H, Gothe, R, Monika,Z. Genetic differentiation of mites of the genus Chorioptes(Acari: Psoroptidae). Exp Appl Acarol,1999,23(4):309-318.
67. Evans, G. O.. Principle of Acarology. CAB International, 1992:1-552.
68. Evans, J.D. Microsatellite loci in the honey bee parasitic mite Varroa jacobsoni. Mol Ecol,2000,9(9) 1436-1438.
69. Evans, W.. Our present knowledge of the fauna of Forth Area. Proc. Phys. Soc, 1909, 17(1): 1-64.
70. H. Bradley Shaffer, Peter Meylan, Mark L. McKnight Tests of Turtle Phylogeny: Molecular, Morphological, and Paleontological Approaches.Systematic Biology, 1997,46(2): 235-268.
71. Harvey M S The Australian water mites: A guide to families and genera. Monographs on Invertebrate Taxonomy. Vol4. CSIRO Publishing, Australia 1998
72. Jeyaprakash A.; Hoy M.A. Mitochondrial 12 S rRNA sequences used to design a molecular ladder assay to identify six commercially available phytoseiids(Acari: Phytoseiidae). Biological Control, 2002, 25(2): 136-142.
73. Krantz, G. W.. A Manual of Acarology(Second Edition). Oregon State Univeristy Bookstore.Inc.Corvallis, 1978:1-221.
74. Lekveishvili, M, Klompen, H. Phylogeny of infraorder Sejina(Acari: Mesostigmata). Zootaxa, 2004,(629): 1-19.
75. Maraun, M, Heethoff, M, Schneider, K, Weigmann, G, Cianciolo, J, Thomas, R, H, Norton, R, A. Molecular phylogeny of oribatid mites(Oribatida, Acari): evidence for multiple radiations of parthenogenetic lineages. Exp Appl Acarol, 2004,33(3): 183-201.
76. Navajas, M.J, Thistlewood, H.M, Lagnel, J, Hughes, C. Microsatellite sequences are under-represented in two mite genomes. Insect Mol Biol,1998,7(3):249-256.
77. Navajas, M, Cotton, D, Kreiter, S. Molecular approach in spider mites(Acari: Tetranychidae): preliminary data on ribosomal DNA sequences. Exp Appl Acarol, 1992,15(4):211-8.
78. Navajas, M, Lagnel, J, Fauvel, G, Moraes, G, DE. Sequence variation of ribosomal internal transcribed spacers(ITS) in commercially important Phytoseiidae mites. Exp Appl Acarol,1999,23(11): 851-859.
79. Nishimura, S, Hinomoto, N, Takafuji, A. Isolation, characterization, inheritance and linkage of microsatellite markers in Tetranychus kanzawai(Acari: Tetranychidae). Exp Appl Acarol, 2003,31(1-2):93-103.
80. Olomski, R.. The evolutionary transformation of osmotic regulation in the life cycle of freshwater mites(Hydracnidae). In Schuster R, Murphy P W ed. The Acari, Reproduction, Development and life-history Strategies. London, Chapman and Hall 57-64.
81. Osakabe, M, Sakagami, Y. RFLP analysis of ribosomal DNA in sibling species of spider mite, genus Panonychus(Acari: Tetranychidae). Insect Mol Biol,1994,3(1):63-66.
82. Perrot-Minno, M.J, Lagnel, J, Desmarais, E, Navajas, M. Isolation and characterization by direct amplification of length polymorphisms(DALP) of codominant genetic markers with Mendelian inheritance in Neoseiulus californicus(Acari: Phytoseiidae). Exp Appl Acarol,2000, 24(10-11):795-803.
83. Pham, X.D, Otsuka, Y, Suzuki, H,. Detection of Orientia tsutsugamushi(Rickettsiales: rickettsiaceae) in unengorged chiggers(Acari: Trombiculidae) from Oita Prefecture, Japan, by nested polymerase chain reaction. J Med Entomol.2001,38(2):308-311.
84. Rey I, Dorda BA, Valdecasas AG. Traditional water mite fixatives and their compatibility with later DNA studies. Exp Appl Acarol JT-Experimental & applied acarology, 2004,34(1-2):59-65.
85. Rodrigues, J.C.V, Gallo-Meagher, M, Ochoa, R, Childers, C.C.C, Adams, B, J. Mitochondrial DNA and RAPD polymorphisms in the haploid mite Brevipalpus phoenicis(Acari: Tenuipalpidae). Exp Appl Acarol,2004,34(3-4): 275-290.
86. Prasad, V.. and Cook, D. R. The taxonomy of water mites larvae. Mem. Amer., Ent. Inst, 1972,(18): 1-326.
87. Salomone, N, Emerson, B.C, Hewitt, G.M, Bernini, F.. Phylogenetic relationships among the Canary Island Steganacaridae(Acari, Oribatida) inferred from mitochondrial DNA sequence data. Mol Ecol,2002,11(1):79-89.
88. Soller, R, Wohltmann, A, Witte, H, Blohm, D.. Phylogenetic relationships within terrestrial mites(Acari: Prostigmata, Parasitengona) inferred from comparative DNA sequence analysis of the mitochondrial cytochrome oxidase subunit I gene. Mol Phylogenet Evol,2001,18(1):47-53.
89. Toda, S, Osakabe, M, Komazaki, S. Interspecific diversity of mitochondrial COI sequences in Japanese Panonychus species(Acari: Tetranychidae). Exp Appl Acarol, 2000, 24(10-11): 821-829.
90. Vargas, M, Garcia-Varela, M, Laclette, J. P, Perez, T. M.. Application of ITS-2 sequences as markers for identification and phylogenetic inference within the genus Geomylichus(Acari: Listrophoridae). Exp Appl Acarol, 2005, 35(3): 223-238.
91. Weeks, A. R, van Opijnen, T, Breeuwer, J. A. AFLP fingerprinting for assessing intraspecific variation and genome mapping in mites. Exp Appl Acarol, 2000, 24(10-11): 775-793.
92. Wiles, P. R.. A new family, genus and species of watermite(Acari: Hydrachnidia, Lebertioidea) from Brunei. Quekandt Journal of Microscopy, 1996, 37: 692-695
93. Witte, H.. Indirect sperm transfer in the prostigmatic mites from a phylogenandic viewpoint. In: Schuster R, Muphy P Wed. The Acari, Reproduction, Development And Life-history Strategies, Chaoman and Hall, 1991. 177-178.
94. Yli-Mattila, T, Paavanen-Huhtala, S, Fenton, B, Tuovinen, T.. Species and strain identification of the predatory mite Euseius finlandicus by RAPD-PCR and ITS sequences. Exp Appl Acarol, 2000, 24(10-11): 863-880.
95. Zahler, M, Essig, A, Gothe, R,. Molecular analyses suggest monospecificity of the genus Sarcoptes(Acari: Sarcoptidae). Int J Pa rasitol, 1999, 29(5): 759-766.
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50. Baker, E. W. and Wharton, G. W.. An Introduction to Acarology. New York: The Macmilan Company, 1952. 1-465.
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61. De Rojas, M, Mora, M.D, Ubeda, J.M, Cutillas.C, Navajas,M, Guevara.D,C. Phylogenetic relationships in rhinonyssid mites(Acari: Rhinonyssidae) based on mitochondrial 16S rDNA sequences. Exp Appl Acarol,2001, 25(12):957-967.
62. De Rojas, M, Mora, M.D, Ubeda, J.M. Phylogenetic relationships in rhinonyssid mites(Acari: Rhinonyssidae) based on ribosomal DNA sequences: Insights for the discrimination of closely related species. Parasitology Research,2002,88(7):675-681.
63. Desmarais, E, Lanneluc, I, Lagnel, J. Direct amplification of length polymorphisms(DALP), or how to get and characterize new genetic markers in many species. Nucleic Acids Res, 1998, 26(6): 1458-1465.
64. Edwardsa, O.R, Melob, E.L, Smith, L, M.A. Hoy. Discrimination of three Typhlodromalus species(Acari: Phytoseiidae) using random amplified polymorphic DNA markers. Exp Appl Acarol,1997, 21: 101-109.
65. Ernsting, B.R, Edwards, D.D, Vidrine, M.F, Myers,K.S, Harmon,C.M. Phylogenetic relationships among species of the subgenus Parasitatax(Acari: Unionicolidae: Unionicola) based on DNA sequence of the mitochondrial cytochrome oxidase I gene. International Journal Of Acarology,2006,32(2): 195-202.
66. Essig, A, Rinder, H, Gothe, R, Monika,Z. Genetic differentiation of mites of the genus Chorioptes(Acari: Psoroptidae). Exp Appl Acarol,1999,23(4):309-318.
67. Evans, G. O.. Principle of Acarology. CAB International, 1992:1-552.
68. Evans, J.D. Microsatellite loci in the honey bee parasitic mite Varroa jacobsoni. Mol Ecol,2000,9(9) 1436-1438.
69. Evans, W.. Our present knowledge of the fauna of Forth Area. Proc. Phys. Soc, 1909, 17(1): 1-64.
70. H. Bradley Shaffer, Peter Meylan, Mark L. McKnight Tests of Turtle Phylogeny: Molecular, Morphological, and Paleontological Approaches.Systematic Biology, 1997,46(2): 235-268.
71. Harvey M S The Australian water mites: A guide to families and genera. Monographs on Invertebrate Taxonomy. Vol4. CSIRO Publishing, Australia 1998
72. Jeyaprakash A.; Hoy M.A. Mitochondrial 12 S rRNA sequences used to design a molecular ladder assay to identify six commercially available phytoseiids(Acari: Phytoseiidae). Biological Control, 2002, 25(2): 136-142.
73. Krantz, G. W.. A Manual of Acarology(Second Edition). Oregon State Univeristy Bookstore.Inc.Corvallis, 1978:1-221.
74. Lekveishvili, M, Klompen, H. Phylogeny of infraorder Sejina(Acari: Mesostigmata). Zootaxa, 2004,(629): 1-19.
75. Maraun, M, Heethoff, M, Schneider, K, Weigmann, G, Cianciolo, J, Thomas, R, H, Norton, R, A. Molecular phylogeny of oribatid mites(Oribatida, Acari): evidence for multiple radiations of parthenogenetic lineages. Exp Appl Acarol, 2004,33(3): 183-201.
76. Navajas, M.J, Thistlewood, H.M, Lagnel, J, Hughes, C. Microsatellite sequences are under-represented in two mite genomes. Insect Mol Biol,1998,7(3):249-256.
77. Navajas, M, Cotton, D, Kreiter, S. Molecular approach in spider mites(Acari: Tetranychidae): preliminary data on ribosomal DNA sequences. Exp Appl Acarol, 1992,15(4):211-8.
78. Navajas, M, Lagnel, J, Fauvel, G, Moraes, G, DE. Sequence variation of ribosomal internal transcribed spacers(ITS) in commercially important Phytoseiidae mites. Exp Appl Acarol,1999,23(11): 851-859.
79. Nishimura, S, Hinomoto, N, Takafuji, A. Isolation, characterization, inheritance and linkage of microsatellite markers in Tetranychus kanzawai(Acari: Tetranychidae). Exp Appl Acarol, 2003,31(1-2):93-103.
80. Olomski, R.. The evolutionary transformation of osmotic regulation in the life cycle of freshwater mites(Hydracnidae). In Schuster R, Murphy P W ed. The Acari, Reproduction, Development and life-history Strategies. London, Chapman and Hall 57-64.
81. Osakabe, M, Sakagami, Y. RFLP analysis of ribosomal DNA in sibling species of spider mite, genus Panonychus(Acari: Tetranychidae). Insect Mol Biol,1994,3(1):63-66.
82. Perrot-Minno, M.J, Lagnel, J, Desmarais, E, Navajas, M. Isolation and characterization by direct amplification of length polymorphisms(DALP) of codominant genetic markers with Mendelian inheritance in Neoseiulus californicus(Acari: Phytoseiidae). Exp Appl Acarol,2000, 24(10-11):795-803.
83. Pham, X.D, Otsuka, Y, Suzuki, H,. Detection of Orientia tsutsugamushi(Rickettsiales: rickettsiaceae) in unengorged chiggers(Acari: Trombiculidae) from Oita Prefecture, Japan, by nested polymerase chain reaction. J Med Entomol.2001,38(2):308-311.
84. Rey I, Dorda BA, Valdecasas AG. Traditional water mite fixatives and their compatibility with later DNA studies. Exp Appl Acarol JT-Experimental & applied acarology, 2004,34(1-2):59-65.
85. Rodrigues, J.C.V, Gallo-Meagher, M, Ochoa, R, Childers, C.C.C, Adams, B, J. Mitochondrial DNA and RAPD polymorphisms in the haploid mite Brevipalpus phoenicis(Acari: Tenuipalpidae). Exp Appl Acarol,2004,34(3-4): 275-290.
86. Prasad, V.. and Cook, D. R. The taxonomy of water mites larvae. Mem. Amer., Ent. Inst, 1972,(18): 1-326.
87. Salomone, N, Emerson, B.C, Hewitt, G.M, Bernini, F.. Phylogenetic relationships among the Canary Island Steganacaridae(Acari, Oribatida) inferred from mitochondrial DNA sequence data. Mol Ecol,2002,11(1):79-89.
88. Soller, R, Wohltmann, A, Witte, H, Blohm, D.. Phylogenetic relationships within terrestrial mites(Acari: Prostigmata, Parasitengona) inferred from comparative DNA sequence analysis of the mitochondrial cytochrome oxidase subunit I gene. Mol Phylogenet Evol,2001,18(1):47-53.
89. Toda, S, Osakabe, M, Komazaki, S. Interspecific diversity of mitochondrial COI sequences in Japanese Panonychus species(Acari: Tetranychidae). Exp Appl Acarol, 2000, 24(10-11): 821-829.
90. Vargas, M, Garcia-Varela, M, Laclette, J. P, Perez, T. M.. Application of ITS-2 sequences as markers for identification and phylogenetic inference within the genus Geomylichus(Acari: Listrophoridae). Exp Appl Acarol, 2005, 35(3): 223-238.
91. Weeks, A. R, van Opijnen, T, Breeuwer, J. A. AFLP fingerprinting for assessing intraspecific variation and genome mapping in mites. Exp Appl Acarol, 2000, 24(10-11): 775-793.
92. Wiles, P. R.. A new family, genus and species of watermite(Acari: Hydrachnidia, Lebertioidea) from Brunei. Quekandt Journal of Microscopy, 1996, 37: 692-695
93. Witte, H.. Indirect sperm transfer in the prostigmatic mites from a phylogenandic viewpoint. In: Schuster R, Muphy P Wed. The Acari, Reproduction, Development And Life-history Strategies, Chaoman and Hall, 1991. 177-178.
94. Yli-Mattila, T, Paavanen-Huhtala, S, Fenton, B, Tuovinen, T.. Species and strain identification of the predatory mite Euseius finlandicus by RAPD-PCR and ITS sequences. Exp Appl Acarol, 2000, 24(10-11): 863-880.
95. Zahler, M, Essig, A, Gothe, R,. Molecular analyses suggest monospecificity of the genus Sarcoptes(Acari: Sarcoptidae). Int J Pa rasitol, 1999, 29(5): 759-766.