须脚目动物系统学研究进展
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摘要
须脚目是一类体型微小、形态特殊、生活于土壤或洞穴中的蛛形动物,目前全世界仅记录109种,隶属于2科8属,包括2个化石记录种。由于对该类群系统学研究比较欠缺,其系统发育地位仍不明确。基于形态学的支序系统学分析大部分支持须脚目与四肺类(蜘蛛目+无鞭目+鞭蝎目+裂盾目)聚为一支,但也有研究支持须脚目与真螨类关系较近;分子系统学方面的研究较少,其分析结果也存在分歧。目前中国须脚目动物仅有零星的报道,研究匮乏。现简要报道最近在广西发现的1种类须脚蝎属动物,分析须脚目已知科、属及其分布,比较世界各地区已知须脚目的物种数,列举须脚目研究学者及其研究区域,从经典分类学和系统发育两个方面对须脚目系统学研究的最新进展进行综述,同时给出了须脚目各科、属的中文学名翻译,以期推动中国须脚目动物系统学的开展。
Palpigradi is a group of tiny, soil or cave dwelling arachnids with special morphology. There are only 109 species of Palpigradi described all over the world, and they are belonging to 2 families and 8 genera, with 2 fossil species recorded up to now. Systematic studies on Palpigradi are relatively deficient and their phylogenetic positions are still under discussion. Most cladistic analyses based on morphological characters support Palpigradi as a sister group of Tetrapulmonata(Araneae+Amblypygi+Thelyphonida+Schizomida), while other studies support a closer relationship between Palpigradi and Acariformes mites. The sparse molecular analyses have similar contradict results. So far, there are very few records of Palpigradi in China. The present article shortly reported a newly found species of genus Koeneniodes and its habitat, analyzed the families and genera of Palpigradi in the world and their distributions, compared the species found in different areas of the world, listed the researchers and their study area.The Chinese name of the families and genera of Palpigradi are also provided. The recent progresses of systematic study of Palpigradi are summarized comprehensively from the aspects of taxonomy and phylogeny in order to drive the study of Palpigradi of China in the future.
Palpigradi is a group of tiny, soil or cave dwelling arachnids with special morphology. There are only 109 species of Palpigradi described all over the world, and they are belonging to 2 families and 8 genera, with 2 fossil species recorded up to now. Systematic studies on Palpigradi are relatively deficient and their phylogenetic positions are still under discussion. Most cladistic analyses based on morphological characters support Palpigradi as a sister group of Tetrapulmonata(Araneae+Amblypygi+Thelyphonida+Schizomida), while other studies support a closer relationship between Palpigradi and Acariformes mites. The sparse molecular analyses have similar contradict results. So far, there are very few records of Palpigradi in China. The present article shortly reported a newly found species of genus Koeneniodes and its habitat, analyzed the families and genera of Palpigradi in the world and their distributions, compared the species found in different areas of the world, listed the researchers and their study area.The Chinese name of the families and genera of Palpigradi are also provided. The recent progresses of systematic study of Palpigradi are summarized comprehensively from the aspects of taxonomy and phylogeny in order to drive the study of Palpigradi of China in the future.
引文
[1]Harvey MS.Catalogue of the smaller arachnid orders of the world:Amblypygi,Uropygi,Schizomida,Palpigradi,Ricinulei and Solifugae[M].Melbourne:CSIROPublishing,2003
[2]Harvey MS.Palpigrades of the World,version 1.0.Western Australian Museum,Perth[EB/OL].(2011).http://museum.wa.gov.au/catalogues-beta/palpigrades.
[3]Krantz GW,Walter DE.A Manual of Acarology[M].3rd edition.Texas:Texas Tech University Press,2009
[4]Pepato AR,da Rocha CE,Dunlop JA.Phylogenetic position of the acariform mites:sensitivity to homology assessment under total evidence.BMC Evol Biol,2010,10:235
[5]Souza MFVR,Ferreira RL.Pandora is on Earth:new species of Eukoenenia(Palpigradi)emerging at risk of extinction.Invertebr Syst,2018,32:581-604
[6]Mayoral JG.Clase Arachnida.Orden Palpigradi[M]//Melic A,Ribera I,Torralba A.Revista IDE@-SEA,2015,10:1-9
[7]Weygoldt P,Paulus HF.Untersuchungen zur Morphologie,Taxonomie und Phylogenie der Chelicerata.II.Cladogramme und die Entfaltung der Chelicerata.Z Zool Syst Evol Forsch,1979,17:177-200
[8]Shultz JW.Evolutionary morphology and phylogeny of Arachnida.Cladistics,1990,6:1-31
[9]Shultz JW.A phylogenetic analysis of the arachnid orders based on morphological characters.Zool J Linn Soc,2007,150:221-65
[10]Giribet G,Edgecombe GD,Wheeler WC,et al.Phylogeny and systematic position of Opiliones:a combined analysis of chelicerate relationships using morphological and molecular data.Cladistics,2002,18:5-70
[11]Giribet G,McIntyre E,Christian E,et al.The first phylogenetic analysis of Palpigradi(Arachnida)--the most enigmatic arthropod order.Invertebr Syst,2014,28:350-60
[12]Regier JC,Shultz JW,Zwick A,et al.Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences.Nature,2010,463:1079-83
[13]CondéB.Palpigrades de Brunei et de Hong Kong.Rev Suisse de Zool,1993,100:25-30
[14]田明义,Deharveng L,Bedos A,等.广西洞穴动物多样性研究进展--以无脊椎动物为主[C].全国第十七届洞穴学术会议论文集,2011:149-63
[15]Grassi B,Calandruccio S.Intorno ad un nuovo Aracnide Artrogastro(Koenenia mirabils[sic])che crediamo rappresentante d’un nuovo ordine(Microteliphonida).Naturalista Siciliano,1885,4:127-33,162-8
[16]Grassi B.I progenitori dei Miriapodi e degliInsetti Memoria V.Intorno ad un nuovo Aracnide artrogastro(Koenenia mirabilis)rappresentante di un nuovo ordine(Microthelyphonida).Boll Soc Entomol Ital,1886,18:153-72
[17]Thorell T.Pedipalpi e Scorpioni dell’Arcipelago Malese conservati nel Museo Civico di Storia Naturale di Genova.Ann Mus Civ Stor Nat Genova,1888,26:327-428
[18]Hansen HJ,S?rensen W.The order Palpigradi Thor.(Koenenia mirabilis Grassi)and its relationship to the other Arachnida.Entomol Tidskrift,1897,18:223-40
[19]Berland L.Arachnides et Crustacès[M]//Perrier R.Faune de la France en tableaux synoptiques.Paris:Librairie Delagrave,1929,2:1-220
[20]Harvey MS.Palpigradi[M]//Walton DW.Zoological catalogue of Australia.Canberra:Australian Government Publishing Service,1985,3:158-9
[21]B?rner C.Arachnologische Studien.IV.Die Genitalorgane der Pedipalpen.Zool Anz,1902,26:81-92
[22]K?stner A.Palpigradi Thorell[M]//Kükenthal W.Handbuch der Zoologie.Berlin und Leipzig:Walter de Gruyter&Co.,1932,3:77-98
[23]Petrunkevitch.Arachnida[M]//Moore RC.Treatise on invertebrate paleontology,Arthropoda.Lawrence:University of Kansas Press,1955,2:42-162
[24]Engel MS,Breitkreuz LCV,Cai C,et al.The first Mesozoic microwhip scorpion(Palpigradi):a new genus and species in mid-Cretaceous amber from Myanmar.Sci Nat,2016,103:19
[25]CondéB.Les Palpigrades,1885-1995:acquisitions et lacunes.Rev Suisse Zool,hors série,1996,1:87-106
[26]Rowland JM,Sissom WD.Report on a fossil palpigrade from the Tertiary of Arizona,and a review of the morphology and systematics of the order(Arachnida).JArachnol,1980,8:69-86
[27]Rémy PA.Palpigrades de Madagascar.Mém Inst Sci Madag,Série A,1950,4:135-64
[28]Rémy PA.Palpigrades de l’?le Maurice.Mauritius Institute Bulletin,1958,5:94-102
[29]Rémy PA.Les Palpigrades de Ceylan et leurécologie.Revue Fran?ais d’Entomologie,1961,28:112-9
[30]CondéB.Palpigrades endogés de Singapour et de l’Indonésie.Rev Suisse Zool,1990,97:681-97
[31]Harvey MS.The Schizomida(Arachnida)of the Seychelle Islands.Invertebr Taxon,2001,15:681-93
[32]Zhang ZQ.Phylum Arthropoda[M]//Zhang ZQ.Animal biodiversity:an outline of higher-level classification and survey of taxonomic richness(Addenda 2013).Zootaxa,2013,3703:1-82
[33]Souza MFVR,Ferreira RL.A new highly troglomorphic species of Eukoenenia(Palpigradi:Eukoeneniidae)from tropical Brazil.J Arachnol,2012,40:151-8
[34]Souza MFVR,Ferreira RL.Eukoenenia virgemdalapa(Palpigradi:Eukoeneniidae):a new troglobitic palpigrade from Brazil.Zootaxa,2012,3295:59-64
[35]Souza MFVR,Ferreira RL.Two new species of the enigmatic Leptokoenenia(Eukoeneniidae:Palpigradi)from Brazil:First Record of the Genus Outside Intertidal Environments.PLoS One,2013,8:e77840
[36]Souza MFVR,Ferreira RL.Two new troglobiotic palpigrades(Palpigradi:Eukoeneniidae)from Brazil.Zootaxa,2016,4171:246-58
[37]Barranco P,Mayoral JG.New palpigrades(Arachnida,Eukoeneniidae)from the Iberian Peninsula.Zootaxa,2014,3826:544-62
[38]Christian E,ChristophoryováJ.Eukoenenia florenciae(Arachnida:Palpigradi):lessons from a newcomer to central Europe and the island of Tenerife.Biologia,2013,6:1182-8
[39]Christian E,Isaia M,Paschetta M,et al.Differentiation among cave populations of the Eukoenenia spelaea species-complex(Arachnida:Palpigradi)in the southwestern Alps.Zootaxa,2014,3794:52-86
[40]Mayoral JG,Barranco P.Rediscovery of the troglobious palpigrade Eukoenenia draco(Peyerimhoff 1906)(Palpigradi:Eukoeneniidae),with notes on the adaptations to a cave-dwelling life.Zootaxa,2013,3635:174-84
[41]Mayoral JG,Barranco P.A remarkable new palpigrade(Arachnida)of the Eukoenenia mirabilis group from Spain.Zootaxa,2017,4290:357-66
[42]Mayoral JG,Barranco P.A cave-dwelling Iberian palpigrade(Arachnida:Palpigradi)of the Eukoenenia mirabilis group.Zootaxa,2017,4363:561-8
[43]Monta?o HM.Redescripción de Eukoenenia hanseni(Arachnida:Palpigradi)y descripción de una nueva especie de palpígrado de México.Rev Iber Aracnol,2012,20:1-15
[44]CondéB.Nouveaux Palpigrades de Trieste,de Slovénie,de Malte,du Paraguay,de Tha?lande et de Bornéo.Rev Suisse de Zool,1988,95:723-50
[45]CondéB.Palpigradida[M]//Juberthie C,Decu,V.Encyclopaedia biospeleologica.Moulis(France)and Bucarest(Roumanei).Sociétéde Biospeologie,1998,2:913-20
[46]CondéB.Palpigrades cavernicoles et endogés de Tha?lande et des Célèbes(1ère note).Rev Suisse de Zool,1992,99:655-72
[47]Harvey MS.The smaller arachnid orders:diversity,descriptions and distributions from Linnaeus to the present(1758 to 2007).Zootaxa,2007,1668:363-80
[48]Ferreira RL,de Oliveira MPA,Souza-Silva M.Biodiversidade subterranea em geossistemas ferruginosos[M]//do Carmo FF,Kamino LHY.Geossistemas Ferruginosos do Brasil.Minas Gerais:Belo Horizonte,2015:195-231
[49]Ferreira RL,Souza MFVR,Machado EO,et al.Description of a new Eukoenenia(Palpigradi:Eukoeneniidae)and Metagonia(Araneae:Pholcidae)from Brazilian caves,with notes on their ecological interactions.J Arachnol,2011,39:409-19
[50]Souza MFVR,Ferreira RL.Eukoenenia(Palpigradi:Eukoeneniidae)in Brazilian caves with the first troglobiotic palpigrade from South America.J Arachnol,2010,38:415-24
[51]Souza MFVR,Ferreira RL.A new troglobitic Eukoenenia(Palpigradi:Eukoeneniidae)from Brazil.J Arachnol,2011,39:185-8
[52]Souza MFVR,Ferreira RL.A new species of Eukoenenia(Palpigradi:Eukoeneniidae)from Brazilian iron ore caves.Zootaxa,2011,2886:31-8
[53]Monta?o HM,Francke OF.Descripción de una nueva especie de palpígrado(Arachnida:Palpigradi)del género Eukoenenia,colectado en la selva lacandona,en México.Entomol Mex,2006,5:162-6
[54]Monta?o HM.Un nuevo palpígrado(Arachnida:Palpigradi)de la Selva Lacandona,México.Rev Iber Aracnol,2007,14:97-103
[55]Wheeler WC,Hayashi CY.The phylogeny of the extant chelicerate orders.Cladistics,1998,14:173-92
[56]van der Hammen L.An introduction to comparative arachnology[M].The Hague:SPB Academic Publishing,1989
[57]Arabi J,Judson ML,Deharveng L,et al.Nucleotide composition of CO1 sequences in Chelicerata(Arthropoda):detecting new mitogenomic rearrangements.J Mol Evol,2012,74:81-95
[58]Sharma PP,Kaluziak ST,Pérez-Porro AR,et al.Phylogenomic interrogation of Arachnida reveals systemic conflicts in phylogenetic signal.Mol Biol Evol,2014,31:2963-84
[2]Harvey MS.Palpigrades of the World,version 1.0.Western Australian Museum,Perth[EB/OL].(2011).http://museum.wa.gov.au/catalogues-beta/palpigrades.
[3]Krantz GW,Walter DE.A Manual of Acarology[M].3rd edition.Texas:Texas Tech University Press,2009
[4]Pepato AR,da Rocha CE,Dunlop JA.Phylogenetic position of the acariform mites:sensitivity to homology assessment under total evidence.BMC Evol Biol,2010,10:235
[5]Souza MFVR,Ferreira RL.Pandora is on Earth:new species of Eukoenenia(Palpigradi)emerging at risk of extinction.Invertebr Syst,2018,32:581-604
[6]Mayoral JG.Clase Arachnida.Orden Palpigradi[M]//Melic A,Ribera I,Torralba A.Revista IDE@-SEA,2015,10:1-9
[7]Weygoldt P,Paulus HF.Untersuchungen zur Morphologie,Taxonomie und Phylogenie der Chelicerata.II.Cladogramme und die Entfaltung der Chelicerata.Z Zool Syst Evol Forsch,1979,17:177-200
[8]Shultz JW.Evolutionary morphology and phylogeny of Arachnida.Cladistics,1990,6:1-31
[9]Shultz JW.A phylogenetic analysis of the arachnid orders based on morphological characters.Zool J Linn Soc,2007,150:221-65
[10]Giribet G,Edgecombe GD,Wheeler WC,et al.Phylogeny and systematic position of Opiliones:a combined analysis of chelicerate relationships using morphological and molecular data.Cladistics,2002,18:5-70
[11]Giribet G,McIntyre E,Christian E,et al.The first phylogenetic analysis of Palpigradi(Arachnida)--the most enigmatic arthropod order.Invertebr Syst,2014,28:350-60
[12]Regier JC,Shultz JW,Zwick A,et al.Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences.Nature,2010,463:1079-83
[13]CondéB.Palpigrades de Brunei et de Hong Kong.Rev Suisse de Zool,1993,100:25-30
[14]田明义,Deharveng L,Bedos A,等.广西洞穴动物多样性研究进展--以无脊椎动物为主[C].全国第十七届洞穴学术会议论文集,2011:149-63
[15]Grassi B,Calandruccio S.Intorno ad un nuovo Aracnide Artrogastro(Koenenia mirabils[sic])che crediamo rappresentante d’un nuovo ordine(Microteliphonida).Naturalista Siciliano,1885,4:127-33,162-8
[16]Grassi B.I progenitori dei Miriapodi e degliInsetti Memoria V.Intorno ad un nuovo Aracnide artrogastro(Koenenia mirabilis)rappresentante di un nuovo ordine(Microthelyphonida).Boll Soc Entomol Ital,1886,18:153-72
[17]Thorell T.Pedipalpi e Scorpioni dell’Arcipelago Malese conservati nel Museo Civico di Storia Naturale di Genova.Ann Mus Civ Stor Nat Genova,1888,26:327-428
[18]Hansen HJ,S?rensen W.The order Palpigradi Thor.(Koenenia mirabilis Grassi)and its relationship to the other Arachnida.Entomol Tidskrift,1897,18:223-40
[19]Berland L.Arachnides et Crustacès[M]//Perrier R.Faune de la France en tableaux synoptiques.Paris:Librairie Delagrave,1929,2:1-220
[20]Harvey MS.Palpigradi[M]//Walton DW.Zoological catalogue of Australia.Canberra:Australian Government Publishing Service,1985,3:158-9
[21]B?rner C.Arachnologische Studien.IV.Die Genitalorgane der Pedipalpen.Zool Anz,1902,26:81-92
[22]K?stner A.Palpigradi Thorell[M]//Kükenthal W.Handbuch der Zoologie.Berlin und Leipzig:Walter de Gruyter&Co.,1932,3:77-98
[23]Petrunkevitch.Arachnida[M]//Moore RC.Treatise on invertebrate paleontology,Arthropoda.Lawrence:University of Kansas Press,1955,2:42-162
[24]Engel MS,Breitkreuz LCV,Cai C,et al.The first Mesozoic microwhip scorpion(Palpigradi):a new genus and species in mid-Cretaceous amber from Myanmar.Sci Nat,2016,103:19
[25]CondéB.Les Palpigrades,1885-1995:acquisitions et lacunes.Rev Suisse Zool,hors série,1996,1:87-106
[26]Rowland JM,Sissom WD.Report on a fossil palpigrade from the Tertiary of Arizona,and a review of the morphology and systematics of the order(Arachnida).JArachnol,1980,8:69-86
[27]Rémy PA.Palpigrades de Madagascar.Mém Inst Sci Madag,Série A,1950,4:135-64
[28]Rémy PA.Palpigrades de l’?le Maurice.Mauritius Institute Bulletin,1958,5:94-102
[29]Rémy PA.Les Palpigrades de Ceylan et leurécologie.Revue Fran?ais d’Entomologie,1961,28:112-9
[30]CondéB.Palpigrades endogés de Singapour et de l’Indonésie.Rev Suisse Zool,1990,97:681-97
[31]Harvey MS.The Schizomida(Arachnida)of the Seychelle Islands.Invertebr Taxon,2001,15:681-93
[32]Zhang ZQ.Phylum Arthropoda[M]//Zhang ZQ.Animal biodiversity:an outline of higher-level classification and survey of taxonomic richness(Addenda 2013).Zootaxa,2013,3703:1-82
[33]Souza MFVR,Ferreira RL.A new highly troglomorphic species of Eukoenenia(Palpigradi:Eukoeneniidae)from tropical Brazil.J Arachnol,2012,40:151-8
[34]Souza MFVR,Ferreira RL.Eukoenenia virgemdalapa(Palpigradi:Eukoeneniidae):a new troglobitic palpigrade from Brazil.Zootaxa,2012,3295:59-64
[35]Souza MFVR,Ferreira RL.Two new species of the enigmatic Leptokoenenia(Eukoeneniidae:Palpigradi)from Brazil:First Record of the Genus Outside Intertidal Environments.PLoS One,2013,8:e77840
[36]Souza MFVR,Ferreira RL.Two new troglobiotic palpigrades(Palpigradi:Eukoeneniidae)from Brazil.Zootaxa,2016,4171:246-58
[37]Barranco P,Mayoral JG.New palpigrades(Arachnida,Eukoeneniidae)from the Iberian Peninsula.Zootaxa,2014,3826:544-62
[38]Christian E,ChristophoryováJ.Eukoenenia florenciae(Arachnida:Palpigradi):lessons from a newcomer to central Europe and the island of Tenerife.Biologia,2013,6:1182-8
[39]Christian E,Isaia M,Paschetta M,et al.Differentiation among cave populations of the Eukoenenia spelaea species-complex(Arachnida:Palpigradi)in the southwestern Alps.Zootaxa,2014,3794:52-86
[40]Mayoral JG,Barranco P.Rediscovery of the troglobious palpigrade Eukoenenia draco(Peyerimhoff 1906)(Palpigradi:Eukoeneniidae),with notes on the adaptations to a cave-dwelling life.Zootaxa,2013,3635:174-84
[41]Mayoral JG,Barranco P.A remarkable new palpigrade(Arachnida)of the Eukoenenia mirabilis group from Spain.Zootaxa,2017,4290:357-66
[42]Mayoral JG,Barranco P.A cave-dwelling Iberian palpigrade(Arachnida:Palpigradi)of the Eukoenenia mirabilis group.Zootaxa,2017,4363:561-8
[43]Monta?o HM.Redescripción de Eukoenenia hanseni(Arachnida:Palpigradi)y descripción de una nueva especie de palpígrado de México.Rev Iber Aracnol,2012,20:1-15
[44]CondéB.Nouveaux Palpigrades de Trieste,de Slovénie,de Malte,du Paraguay,de Tha?lande et de Bornéo.Rev Suisse de Zool,1988,95:723-50
[45]CondéB.Palpigradida[M]//Juberthie C,Decu,V.Encyclopaedia biospeleologica.Moulis(France)and Bucarest(Roumanei).Sociétéde Biospeologie,1998,2:913-20
[46]CondéB.Palpigrades cavernicoles et endogés de Tha?lande et des Célèbes(1ère note).Rev Suisse de Zool,1992,99:655-72
[47]Harvey MS.The smaller arachnid orders:diversity,descriptions and distributions from Linnaeus to the present(1758 to 2007).Zootaxa,2007,1668:363-80
[48]Ferreira RL,de Oliveira MPA,Souza-Silva M.Biodiversidade subterranea em geossistemas ferruginosos[M]//do Carmo FF,Kamino LHY.Geossistemas Ferruginosos do Brasil.Minas Gerais:Belo Horizonte,2015:195-231
[49]Ferreira RL,Souza MFVR,Machado EO,et al.Description of a new Eukoenenia(Palpigradi:Eukoeneniidae)and Metagonia(Araneae:Pholcidae)from Brazilian caves,with notes on their ecological interactions.J Arachnol,2011,39:409-19
[50]Souza MFVR,Ferreira RL.Eukoenenia(Palpigradi:Eukoeneniidae)in Brazilian caves with the first troglobiotic palpigrade from South America.J Arachnol,2010,38:415-24
[51]Souza MFVR,Ferreira RL.A new troglobitic Eukoenenia(Palpigradi:Eukoeneniidae)from Brazil.J Arachnol,2011,39:185-8
[52]Souza MFVR,Ferreira RL.A new species of Eukoenenia(Palpigradi:Eukoeneniidae)from Brazilian iron ore caves.Zootaxa,2011,2886:31-8
[53]Monta?o HM,Francke OF.Descripción de una nueva especie de palpígrado(Arachnida:Palpigradi)del género Eukoenenia,colectado en la selva lacandona,en México.Entomol Mex,2006,5:162-6
[54]Monta?o HM.Un nuevo palpígrado(Arachnida:Palpigradi)de la Selva Lacandona,México.Rev Iber Aracnol,2007,14:97-103
[55]Wheeler WC,Hayashi CY.The phylogeny of the extant chelicerate orders.Cladistics,1998,14:173-92
[56]van der Hammen L.An introduction to comparative arachnology[M].The Hague:SPB Academic Publishing,1989
[57]Arabi J,Judson ML,Deharveng L,et al.Nucleotide composition of CO1 sequences in Chelicerata(Arthropoda):detecting new mitogenomic rearrangements.J Mol Evol,2012,74:81-95
[58]Sharma PP,Kaluziak ST,Pérez-Porro AR,et al.Phylogenomic interrogation of Arachnida reveals systemic conflicts in phylogenetic signal.Mol Biol Evol,2014,31:2963-84
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