云南蜘蛛?yún)^(qū)系初步分析*

楊自忠，方 波，李成功，何 苗，李 玥

(1.大理大學(xué)云南省昆蟲醫(yī)藥研發(fā)重點實驗室，云南 大理 671000 ; 2.云南省林業(yè)科學(xué)院,云南 昆明 650001)

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云南蜘蛛?yún)^(qū)系初步分析*

楊自忠1，方 波2，李成功1，何 苗1，李 玥1

(1.大理大學(xué)云南省昆蟲醫(yī)藥研發(fā)重點實驗室，云南 大理 671000 ; 2.云南省林業(yè)科學(xué)院,云南 昆明 650001)

本文通過整理多年來對云南蜘蛛?yún)^(qū)系的研究結(jié)果，結(jié)合國內(nèi)外學(xué)者的研究報道，系統(tǒng)報道了云南蜘蛛58科389屬1248種，其中，云南特有物種665種。同時，結(jié)合物種分布特點和地質(zhì)歷史資料對云南蜘蛛?yún)^(qū)系的特點做了簡單的分析，分析結(jié)果表明云南蜘蛛?yún)^(qū)系特點為：物種多樣性(diversity)復(fù)雜；特有種(Endemic species)豐富；區(qū)系古老，既有屬于岡瓦納古陸(Gondwanaland)起源的舊熱帶區(qū)系成分，也有勞亞古陸(Lanrasia)起源的北溫帶區(qū)系成分。云南蜘蛛?yún)^(qū)系勞亞古陸和岡瓦納古陸區(qū)系成分相互滲透以及一些滲透進來的古老物種因適應(yīng)云南特殊地理環(huán)境和氣候環(huán)境演化發(fā)展的結(jié)果。

蜘蛛?yún)^(qū)系(Fauna of Spider)；特有種(Endemic species)；岡瓦納古陸(Gondwanaland)；勞亞古陸(Lanrasia)；云南

蜘蛛隸屬于節(jié)肢動物門(Arthropoda)蛛形綱(Arachnoidea)蜘蛛目(Araneae)，起源于古生代(Paleozoic)前泥盆紀(jì)(Devonian)；是現(xiàn)在陸地上多樣性最豐富的捕食者類群之一。通常，蜘蛛(spider)主要以獵捕活體昆蟲為食，少數(shù)獵食如孔蛛、花皮蛛等其他蜘蛛，大型種類還能捕食其他小型動物，如青蛙等。通過幾億年的演化，蜘蛛由單一的地面類群發(fā)展成今天的結(jié)網(wǎng)和游獵兩大類群，占據(jù)了土壤、落葉、灌叢、森林樹木及水域等等各種生態(tài)空間，其豐富多樣的種類、獨特的生活方式和廣闊的分布空間成了森林和農(nóng)田害蟲的主要天敵。成為維持自然生態(tài)平衡重要的生態(tài)因素。

云南為低緯度高原地區(qū)，境內(nèi)地貌復(fù)雜，氣候多樣，物種生物多樣性豐富，素有“動植物王國”之稱，其中蜘蛛物種多樣性也不例外。對于云南蜘蛛的報道，在上個世紀(jì)只見一些零星的新種報道[1-112]，通過多年的標(biāo)本采集和分類研究，結(jié)合世界蜘蛛名錄及國內(nèi)外學(xué)者研究報道文獻整理出云南蜘蛛共計59科389屬1248種，其中，云南特有物種665種。同時，根據(jù)物種的種屬分布情況，對云南蜘蛛?yún)^(qū)系特點做了初步的分析，現(xiàn)報道如下：

1.物種組成

根據(jù)筆者的調(diào)查研究及對世界蜘蛛名錄(World Spider Catalog Version 16.5)中記錄的整理，共報道云南蜘蛛共計58科389屬1248種[131]，如表1。

表1 云南蜘蛛物種多樣性統(tǒng)計

2.區(qū)系特點(Characteristics of spider fauna)

通過整體分析，云南蜘蛛?yún)^(qū)系具有物種多樣性復(fù)雜、特有種(Endemic species)豐富、區(qū)系古老，既有屬于岡瓦納古陸(Gondwanaland)起源的舊熱帶區(qū)系成分，也有勞亞古陸(Lanrasia)起源的北溫帶區(qū)系成分的特點。

2.1物種多樣性復(fù)雜

根據(jù)近幾年的調(diào)查和現(xiàn)有資料統(tǒng)計，目前已發(fā)現(xiàn)分布于云南的蜘蛛有59科389屬1248種，分別占我國已記述種類的85.50%、52.78%、29.10%，科級階元豐富度極為突出，屬級階元次之，種級階元稍低，基本符合云南蜘蛛物種豐富的特點。種級階元稍低可能與跟調(diào)查不夠相關(guān)。

表1統(tǒng)計顯示：屬級階元中，Salticidae、Linyphiidae、Thomisidae、Theridiidae、Araneidae是云南蜘蛛?yún)^(qū)系的主要類群，分別為53屬、43屬、40屬、39屬和27屬，分別占云南蜘蛛?yún)^(qū)系屬級階元的13.62%，11.05%，10.28%，10.02%和6.94%；而種級階元中，物種比較豐富的依次為Salticidae、Theridiidae、Araneidae和Thomisidae，物種數(shù)量分別為142種、135種、130種和110種，分別占云南蜘蛛?yún)^(qū)系的11.37%，10.81%，10.41%和8.81%，物種豐富度與屬級階元的豐富度吻合。種級階元中，Linyphiidae的物種72種，這與Linyphiidae是蜘蛛類群中物種多樣性最豐富的類群不一致，從中反映出對Linyphiidae調(diào)查還不夠。從已經(jīng)統(tǒng)計得到的數(shù)據(jù)看出，Ctenizidae、Anyphaenidae、Deinopidae、Desidae、Eresidae、Selenopidae和Sinopimoidae僅有1屬1種，盡管種類數(shù)量少，卻是該地區(qū)的蜘蛛?yún)^(qū)系豐富多彩的主要因素。從科、屬種的數(shù)量來看，云南蜘蛛?yún)^(qū)系分別占我國科、屬、種的85.50%、52.78%、29.10%。由此可見，云南的蜘蛛?yún)^(qū)系具有科、屬較豐富，種類相對較少的特點。

2.2特有種(Endemic species)豐富

指目前僅發(fā)現(xiàn)于云南的類群，是云南蜘蛛?yún)^(qū)系主要的成分之一，有665種，占區(qū)系成分的53.28%，特有種占了云南蜘蛛物種的一半以上，這從另外一個角度也充分反映了云南蜘蛛物種的豐富性。在已知的665種特有種中，主要集中分布在滇南的西雙版納、思茅地區(qū)和滇西的大理、保山、德宏地區(qū)、滇西北的怒江、迪慶的高山峽谷地區(qū)。這種分布不均勻特點也反映出了云南南部和西北部云南動物區(qū)系(Fauna)的兩大分化地區(qū)。

3.區(qū)系古老，既有屬于岡瓦納古陸(Gondwanaland)起源的舊熱帶區(qū)系成分，也有勞亞古陸(Lanrasia)起源的北溫帶區(qū)系成分

地史資料證明，二疊紀(jì)末期聯(lián)合古陸形成，中國內(nèi)地的主體既不屬于勞亞古陸，也不屬于岡瓦納古陸，而是介于這兩大古陸之間的特提斯(Tethys)海洋盆系統(tǒng)中的塊體[132]。中生代晚期，岡瓦納古陸開始斷裂解體為非洲-南美洲和印度-澳大利亞-南極洲板塊，隨后印度板塊與澳大利亞-南極洲板塊分離，并向北移動。伴隨著印度板塊的不斷前進，古特提斯海也在逐漸縮小，最終約在5000萬年前與歐亞大陸板塊縫合。在岡瓦納古陸發(fā)生主要斷裂的同時，原澳大利亞-新幾內(nèi)亞板塊北緣的一些小型陸塊(可能包括今天西藏的南部、緬甸、馬來西亞、蘇門答臘島等)也先后斷裂并向東北漂移，最終與歐亞板塊南端拼合[133]。

云南的地史資料表明，古生代期，川滇古陸即隆起形成，陸地范圍雖經(jīng)多次變化，但從未被海水全部淹沒[134]，經(jīng)過中生代三疊紀(jì)的抬升，到了第三紀(jì)末期，云、貴、川完全露出海面形成川滇黔準(zhǔn)平原。云南陸地的發(fā)展歷史，也就是云南蜘蛛?yún)^(qū)的發(fā)展歷史。由此可見，云南蜘蛛?yún)^(qū)系具有悠久的發(fā)展歷史。

3.1區(qū)系古老，多古老蜘蛛的后裔

在蜘蛛目中，中紡亞目的節(jié)板蛛科蜘蛛被公認(rèn)為是最古老的蜘蛛。該類群全球已記述8屬96種，我國有5屬23種，云南報道了1屬2種：云南七紡蛛Heptathela yunnanensis Song & Haupt, 1984分布于滇中的昆明地區(qū)，而峨山七紡蛛H. wosanensis Wang & Jiao, 1995發(fā)現(xiàn)于滇中的峨山地區(qū)，目前又在大理蒼山和賓川的雞足山也發(fā)現(xiàn)了七紡蛛蹤影。

在后紡亞目中，原蛛類是比較低等原始的蜘蛛，全球已記述的有15科329屬2786種，我國有6科17屬68種，云南目前發(fā)現(xiàn)5科5屬13種，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。在科級階元中，云南的原始低等的蜘蛛類群占世界原蛛的33.33%，占我國原蛛類的83.33%。在新蛛下目的蜘蛛類群中，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具屬于勞亞古陸起源的北溫帶區(qū)系成分

根據(jù)云南的地質(zhì)歷史和上述云南蜘蛛名錄及其分布資料可以看出，一些廣泛分布北方(勞亞古陸)的類群，在云南也有分布。在北方廣泛分布的類群，如Tmeticus，在我國除云南有分布外，其他地區(qū)尚未發(fā)現(xiàn)，而典型的北方類群Alopecosa、Talavera、Walckenaeria廣泛分布于我國的新疆、西藏、吉林、遼寧、甘肅、陜西、河北、北京，在云南最南端分布到了滇西的大理和滇東北的永善、昭通地區(qū)；而Araniella、Sitticus、Micaria廣泛分布在于我國的內(nèi)蒙古、陜西、甘肅、青海、河北，最南端沒有越過河北，而在云南抵達了滇西的大理、滇東北的永善；Drassyllus廣泛分布于我國的遼寧、北京、甘肅、新疆、西藏、陜西、河南，偶見于南方的安徽、浙江，在云南分布到了昆明，最南抵達了緬甸；Xysticus除了在我國各地都有分布，在云南也廣泛分布。從總體上分析，北方的區(qū)系成分在云南的分布多數(shù)南伸到了滇西北和滇東北地區(qū)，最南延伸到了昆明。這些屬于勞亞古陸起源的溫帶區(qū)系成分在云南的廣泛分布，說明了云南蜘蛛?yún)^(qū)系與勞亞古陸起源的溫帶區(qū)系成分具有一定的歷史淵源。

3.3具岡瓦納古陸起源的舊熱帶區(qū)系成分

根據(jù)上述云南蜘蛛名錄及其分布資料同樣可以看出，一些廣泛分布于南大陸(岡瓦納古陸)的類群在云南也有分布。如廣泛分布于熱帶非洲區(qū)的Scytodes、Micropholcus、Theridula、Nesticodes、Selenops，熱帶非洲、南美洲和澳洲共有類群Ariadna、Tmarus，非洲和澳洲及東南亞共有類群Nephila、Gea、Gasteracantha、Cyrtarachne、Cyrtophora、Dolomedes、Mallinella、Olios、Hasarius、Myrmarachne，非洲和東南亞地區(qū)共有類群Spermophora、Stegodyphus、Argyrodes、Caerostris、Neoscona、Paraplectana、Thelacantha、Perenethis、Thalassius、Agelena、Stiphropus、Phrynarachne、Monaeses、Runcinia、Thomisus、Carrhotus、Hyllus、Langona、Rhene，非洲、南美洲、澳洲和東南亞地區(qū)共有類群Deinopis、Trochosa，非洲、澳洲和東南亞地區(qū)共有類群有Miagrammopes、Oxyopes，非洲和東南亞地區(qū)共有類群Philoponella、Chrysso、Craspedisia、Mangora、Parawixia、Peucetia，澳洲和東南亞地區(qū)共有類群Mesida、Orsinome、Heteropoda、Telamonia，非洲、南美洲和東南亞地區(qū)共有類群Leucauge和廣泛分布再澳洲的Diaea、Badumna，廣泛分布于非洲的Cyrba、Heliophanus、Menemerus、Thyene、Thyenula，廣泛分布于印度Hippasa等。這些類群在云南都有不同數(shù)量的種類分布。這種分布特點反映了云南蜘蛛?yún)^(qū)系與非洲、南美洲、澳洲及印度次大陸區(qū)系的親緣性。

綜上所述，現(xiàn)在的云南蜘蛛?yún)^(qū)系是勞亞古陸和岡瓦納古陸區(qū)系成分相互滲透以及一些滲透進來的古老物種因適應(yīng)云南特殊地理環(huán)境和氣候環(huán)境演化發(fā)展的結(jié)果。

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(責(zé)任編輯 徐成東)

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

國家自然科學(xué)基金資助項目，項目編號：31060070；中國博士后科學(xué)基金項目，項目編號：20090450677；云南省科技廳應(yīng)用基礎(chǔ)研究面上項目，項目編號：2009CD092；云南省教育廳重點基金項目，項目編號：2011Z107。

2016 - 05 - 13

楊自忠(1969—)，博士，教授，碩士生導(dǎo)師，從事蜘蛛?yún)^(qū)系分類及生態(tài)研究。

Q959.226.2

A

1671 - 7406(2016)06 - 0015 - 11