杨自忠,方 波,李成功,何 苗,李 玥
(1.大理大学云南省昆虫医药研发重点实验室,云南 大理 671000 ; 2.云南省林业科学院,云南 昆明 650001)
云南蜘蛛区系初步分析*
杨自忠1,方 波2,李成功1,何 苗1,李 玥1
(1.大理大学云南省昆虫医药研发重点实验室,云南 大理 671000 ; 2.云南省林业科学院,云南 昆明 650001)
本文通过整理多年来对云南蜘蛛区系的研究结果,结合国内外学者的研究报道,系统报道了云南蜘蛛58科389属1248种,其中,云南特有物种665种。同时,结合物种分布特点和地质历史资料对云南蜘蛛区系的特点做了简单的分析,分析结果表明云南蜘蛛区系特点为:物种多样性(diversity)复杂;特有种(Endemic species)丰富;区系古老,既有属于冈瓦纳古陆(Gondwanaland)起源的旧热带区系成分,也有劳亚古陆(Lanrasia)起源的北温带区系成分。云南蜘蛛区系劳亚古陆和冈瓦纳古陆区系成分相互渗透以及一些渗透进来的古老物种因适应云南特殊地理环境和气候环境演化发展的结果。
蜘蛛区系(Fauna of Spider);特有种(Endemic species);冈瓦纳古陆(Gondwanaland);劳亚古陆(Lanrasia);云南
蜘蛛隶属于节肢动物门(Arthropoda)蛛形纲(Arachnoidea)蜘蛛目(Araneae),起源于古生代(Paleozoic)前泥盆纪(Devonian);是现在陆地上多样性最丰富的捕食者类群之一。通常,蜘蛛(spider)主要以猎捕活体昆虫为食,少数猎食如孔蛛、花皮蛛等其他蜘蛛,大型种类还能捕食其他小型动物,如青蛙等。通过几亿年的演化,蜘蛛由单一的地面类群发展成今天的结网和游猎两大类群,占据了土壤、落叶、灌丛、森林树木及水域等等各种生态空间,其丰富多样的种类、独特的生活方式和广阔的分布空间成了森林和农田害虫的主要天敌。成为维持自然生态平衡重要的生态因素。
云南为低纬度高原地区,境内地貌复杂,气候多样,物种生物多样性丰富,素有“动植物王国”之称,其中蜘蛛物种多样性也不例外。对于云南蜘蛛的报道,在上个世纪只见一些零星的新种报道[1-112],通过多年的标本采集和分类研究,结合世界蜘蛛名录及国内外学者研究报道文献整理出云南蜘蛛共计59科389属1248种,其中,云南特有物种665种。同时,根据物种的种属分布情况,对云南蜘蛛区系特点做了初步的分析,现报道如下:
根据笔者的调查研究及对世界蜘蛛名录(World Spider Catalog Version 16.5)中记录的整理,共报道云南蜘蛛共计58科389属1248种[131],如表1。
表1 云南蜘蛛物种多样性统计
通过整体分析,云南蜘蛛区系具有物种多样性复杂、特有种(Endemic species)丰富、区系古老,既有属于冈瓦纳古陆(Gondwanaland)起源的旧热带区系成分,也有劳亚古陆(Lanrasia)起源的北温带区系成分的特点。
2.1物种多样性复杂
根据近几年的调查和现有资料统计,目前已发现分布于云南的蜘蛛有59科389属1248种,分别占我国已记述种类的85.50%、52.78%、29.10%,科级阶元丰富度极为突出,属级阶元次之,种级阶元稍低,基本符合云南蜘蛛物种丰富的特点。种级阶元稍低可能与跟调查不够相关。
表1统计显示:属级阶元中,Salticidae、Linyphiidae、Thomisidae、Theridiidae、Araneidae是云南蜘蛛区系的主要类群,分别为53属、43属、40属、39属和27属,分别占云南蜘蛛区系属级阶元的13.62%,11.05%,10.28%,10.02%和6.94%;而种级阶元中,物种比较丰富的依次为Salticidae、Theridiidae、Araneidae和Thomisidae,物种数量分别为142种、135种、130种和110种,分别占云南蜘蛛区系的11.37%,10.81%,10.41%和8.81%,物种丰富度与属级阶元的丰富度吻合。种级阶元中,Linyphiidae的物种72种,这与Linyphiidae是蜘蛛类群中物种多样性最丰富的类群不一致,从中反映出对Linyphiidae调查还不够。从已经统计得到的数据看出,Ctenizidae、Anyphaenidae、Deinopidae、Desidae、Eresidae、Selenopidae和Sinopimoidae仅有1属1种,尽管种类数量少,却是该地区的蜘蛛区系丰富多彩的主要因素。从科、属种的数量来看,云南蜘蛛区系分别占我国科、属、种的85.50%、52.78%、29.10%。由此可见,云南的蜘蛛区系具有科、属较丰富,种类相对较少的特点。
2.2特有种(Endemic species)丰富
指目前仅发现于云南的类群,是云南蜘蛛区系主要的成分之一,有665种,占区系成分的53.28%,特有种占了云南蜘蛛物种的一半以上,这从另外一个角度也充分反映了云南蜘蛛物种的丰富性。在已知的665种特有种中,主要集中分布在滇南的西双版纳、思茅地区和滇西的大理、保山、德宏地区、滇西北的怒江、迪庆的高山峡谷地区。这种分布不均匀特点也反映出了云南南部和西北部云南动物区系(Fauna)的两大分化地区。
地史资料证明,二叠纪末期联合古陆形成,中国内地的主体既不属于劳亚古陆,也不属于冈瓦纳古陆,而是介于这两大古陆之间的特提斯(Tethys)海洋盆系统中的块体[132]。中生代晚期,冈瓦纳古陆开始断裂解体为非洲-南美洲和印度-澳大利亚-南极洲板块,随后印度板块与澳大利亚-南极洲板块分离,并向北移动。伴随着印度板块的不断前进,古特提斯海也在逐渐缩小,最终约在5000万年前与欧亚大陆板块缝合。在冈瓦纳古陆发生主要断裂的同时,原澳大利亚-新几内亚板块北缘的一些小型陆块(可能包括今天西藏的南部、缅甸、马来西亚、苏门答腊岛等)也先后断裂并向东北漂移,最终与欧亚板块南端拼合[133]。
云南的地史资料表明,古生代期,川滇古陆即隆起形成,陆地范围虽经多次变化,但从未被海水全部淹没[134],经过中生代三叠纪的抬升,到了第三纪末期,云、贵、川完全露出海面形成川滇黔准平原。云南陆地的发展历史,也就是云南蜘蛛区的发展历史。由此可见,云南蜘蛛区系具有悠久的发展历史。
3.1区系古老,多古老蜘蛛的后裔
在蜘蛛目中,中纺亚目的节板蛛科蜘蛛被公认为是最古老的蜘蛛。该类群全球已记述8属96种,我国有5属23种,云南报道了1属2种:云南七纺蛛Heptathela yunnanensis Song & Haupt, 1984分布于滇中的昆明地区,而峨山七纺蛛H. wosanensis Wang & Jiao, 1995发现于滇中的峨山地区,目前又在大理苍山和宾川的鸡足山也发现了七纺蛛踪影。
在后纺亚目中,原蛛类是比较低等原始的蜘蛛,全球已记述的有15科329属2786种,我国有6科17属68种,云南目前发现5科5属13种,如分布于滇南、滇西南的MacrotheleyunnanicaZhu & Song, 2000、M.bannaensisXu & Yin, 2001、CyclocosmialannaensisSchwendinger, 2005等,而分布于滇西和西北的有M.yaniXu et al., 2002、AtypuspedicellatusZhuetal., 2006、A.sacculatusZhuetal., 2006等。在科级阶元中,云南的原始低等的蜘蛛类群占世界原蛛的33.33%,占我国原蛛类的83.33%。在新蛛下目的蜘蛛类群中,也不乏古老蜘蛛的后裔,如DeinopisliukuensisYin, Griswold et Yan, 2002、PimoaanatolicaHormiga, 1994、P.lihengaeGriswold, Long & Hormiga, 1999、P.reniformisXu & Li, 2007、PutaoahuapingHormiga & Tu, 2008和WeintrauboayunnanYang, Zhu & Song, 2006等。
3.2具属于劳亚古陆起源的北温带区系成分
根据云南的地质历史和上述云南蜘蛛名录及其分布资料可以看出,一些广泛分布北方(劳亚古陆)的类群,在云南也有分布。在北方广泛分布的类群,如Tmeticus,在我国除云南有分布外,其他地区尚未发现,而典型的北方类群Alopecosa、Talavera、Walckenaeria广泛分布于我国的新疆、西藏、吉林、辽宁、甘肃、陕西、河北、北京,在云南最南端分布到了滇西的大理和滇东北的永善、昭通地区;而Araniella、Sitticus、Micaria广泛分布在于我国的内蒙古、陕西、甘肃、青海、河北,最南端没有越过河北,而在云南抵达了滇西的大理、滇东北的永善;Drassyllus广泛分布于我国的辽宁、北京、甘肃、新疆、西藏、陕西、河南,偶见于南方的安徽、浙江,在云南分布到了昆明,最南抵达了缅甸;Xysticus除了在我国各地都有分布,在云南也广泛分布。从总体上分析,北方的区系成分在云南的分布多数南伸到了滇西北和滇东北地区,最南延伸到了昆明。这些属于劳亚古陆起源的温带区系成分在云南的广泛分布,说明了云南蜘蛛区系与劳亚古陆起源的温带区系成分具有一定的历史渊源。
3.3具冈瓦纳古陆起源的旧热带区系成分
根据上述云南蜘蛛名录及其分布资料同样可以看出,一些广泛分布于南大陆(冈瓦纳古陆)的类群在云南也有分布。如广泛分布于热带非洲区的Scytodes、Micropholcus、Theridula、Nesticodes、Selenops,热带非洲、南美洲和澳洲共有类群Ariadna、Tmarus,非洲和澳洲及东南亚共有类群Nephila、Gea、Gasteracantha、Cyrtarachne、Cyrtophora、Dolomedes、Mallinella、Olios、Hasarius、Myrmarachne,非洲和东南亚地区共有类群Spermophora、Stegodyphus、Argyrodes、Caerostris、Neoscona、Paraplectana、Thelacantha、Perenethis、Thalassius、Agelena、Stiphropus、Phrynarachne、Monaeses、Runcinia、Thomisus、Carrhotus、Hyllus、Langona、Rhene,非洲、南美洲、澳洲和东南亚地区共有类群Deinopis、Trochosa,非洲、澳洲和东南亚地区共有类群有Miagrammopes、Oxyopes,非洲和东南亚地区共有类群Philoponella、Chrysso、Craspedisia、Mangora、Parawixia、Peucetia,澳洲和东南亚地区共有类群Mesida、Orsinome、Heteropoda、Telamonia,非洲、南美洲和东南亚地区共有类群Leucauge和广泛分布再澳洲的Diaea、Badumna,广泛分布于非洲的Cyrba、Heliophanus、Menemerus、Thyene、Thyenula,广泛分布于印度Hippasa等。这些类群在云南都有不同数量的种类分布。这种分布特点反映了云南蜘蛛区系与非洲、南美洲、澳洲及印度次大陆区系的亲缘性。
综上所述,现在的云南蜘蛛区系是劳亚古陆和冈瓦纳古陆区系成分相互渗透以及一些渗透进来的古老物种因适应云南特殊地理环境和气候环境演化发展的结果。
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(责任编辑 徐成东)
A Preliminary Analysis of Spider Fauna in Yunnan
YANG Zizhong, FANG Bo, LI Chenggong, HE Miao & LI Yue
(KeyLaboratoryofEntomoceuticsBiopharmaceuticalofYunnan,DaliUniversity,Dali, 671000,YunnanProvince;YunnanAcademyofForestry,Kunming, 650001,YunnanProvince)
Through summarizing years of study results on spider fauna of Yunnan and combining study reports from domestic and international researchers, 1,248 Yunnan spider species belonging to 389 genera of 59 families were systematically reported in this article, including 665 endemic species in Yunnan. Meanwhile, features of spider fauna in Yunnnan were preliminarily analyzed based on species distribution characteristics and geological history data. According to the analysis results, the spider fauna features of Yunnan are: complex species diversity, abundant endemic species, ancient fauna including both paleotropical faunal elements originated from Gondwana and north temperate zone faunal elements originated from Laurasia. The current Yunnan spider fauna result from the interpenetration of spider faunal elements originated from Gondwana and Laurasia, as well as the evolution and development of some penetrated ancient species oriented by adaptation to the special geographical and climatic environment of Yunnan.
Spider fauna; Endemic species; Gondwana; Lanrasia; Yunnan
国家自然科学基金资助项目,项目编号:31060070;中国博士后科学基金项目,项目编号:20090450677;云南省科技厅应用基础研究面上项目,项目编号:2009CD092;云南省教育厅重点基金项目,项目编号:2011Z107。
2016 - 05 - 13
杨自忠(1969—),博士,教授,硕士生导师,从事蜘蛛区系分类及生态研究。
Q959.226.2
A
1671 - 7406(2016)06 - 0015 - 11