云南古猿系统分类研究新进展
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摘要
已在云南4个地点发现晚中新世古猿化石:开远小龙潭,保山羊邑,禄丰石灰坝,元谋小河、竹棚、雷老。石灰坝和元谋是欧亚大陆发现古猿化石最丰富的地点,这些古猿蕴涵着重建晚新生代古猿进化史、生物地理和古生物学的重要信息。本文侧重介绍云南古猿的分类和命名,并对其系统关系和生物地理作初步分析。通过古猿化石形态模式及其变异程度的观察,初步认为:各个地点古猿化石分别代表了一个具有性二型的单一种。暂时认为禄丰石灰坝、开远小龙潭和元谋的古猿化石分别属于一个属内的两个种,其有效学名分别为Lufengpithecuslufen gensis(石灰坝标本)和L keiyuanensis(小龙潭和元谋标本)。从系统学观点看,有证据显示禄丰古猿属既为代表猩猩和人亚科共同姐妹单元原始的人科,也是代表猩猩亚科的原始姊妹单元,我们更趋向于赞同后一选择,但承认还需要通过进一步的综合对比研究来证实。迄今云南发现的古猿化石对研究人猿超科的进化历史提供独特的前景:晚中新世~上新世(8MaBP 2MaBP)时期,当人猿超科在欧亚大陆其它地区都已灭绝时,它们却连续地在云南出现。这表明中国南方或东南亚地区是人猿超科(包括猩猩和长臂猿的祖先)的一个重要的避难所。青藏高原的隆升及其对区域环境的影响,对古猿所造成的生态和地理隔离,可能是古猿?
Late Miocene and Pliocene hominoids have been discovered in four sites or site (complexes): Xiaolongtan, Shihuiba, Yuanmou and Yangyi of Yunnan Province, South China. Shihuiba and Yuanmou are sites with the most prolific hominoid fossil in Eurasia, which have yielded important critical evidence for documenting the evolution history, biogeography and paleobiology of Late Neogene hominoids. The aim of this paper is to clarify their taxonomy and nomenclature and present a preliminary synthesis of their phylogenetic relationship and biogeography. According to the morphological pattern and variation degree of the fossil samples, the hominoid fossils in each site respectively represent a single, sexually dimorohic species. Provisionally, we consider the Shihuiba, Xiaolongtan and Yuanmou samples to be two separate species of a single genus. The valid names for these species are Lufengpithecus lufengensis(from Shihuiba)and L. keiyuanensis(from Xiaolongtan and Yuanmou). From a phylogenetic perspective, the present available evidence suggests that lufengensis is either a primitive hominid that represents the sister taxon of the Ponginae + Homininae or a primitive sister taxon to the Ponginae. We tend to favor the second alternative, but acknowledge that a more comprehensive comparative analysis is needed to substantiate the phylogenetic and taxonomic affinities of Lufengpithcus. Importantly, the fossil apes of Yunnan provide a unique temporal perspective in the study on the evolution history of hominoeds. Their continuous occurrence during the Late Miocene and Pliocene(8Ma~2Ma), when hominoids became extinct throughout the rest of Eurasia, suggests that South China(and presumably Southeast Asia in general)was an important refugium for hominoids, including the ancestors of the orang-utans and gibbons. The uplift of the Tibetan plateau and its impact on regional climatic conditions may be an important factor in isolating the hominoids geographically and ecologically. We speculate that the changed climatic condition in the Mid-Pliocene and possibly the arrival of Homo soon after may have precipitated the regional extinction of large hominoids in South China and Southeast Asia mainland.
Late Miocene and Pliocene hominoids have been discovered in four sites or site (complexes): Xiaolongtan, Shihuiba, Yuanmou and Yangyi of Yunnan Province, South China. Shihuiba and Yuanmou are sites with the most prolific hominoid fossil in Eurasia, which have yielded important critical evidence for documenting the evolution history, biogeography and paleobiology of Late Neogene hominoids. The aim of this paper is to clarify their taxonomy and nomenclature and present a preliminary synthesis of their phylogenetic relationship and biogeography. According to the morphological pattern and variation degree of the fossil samples, the hominoid fossils in each site respectively represent a single, sexually dimorohic species. Provisionally, we consider the Shihuiba, Xiaolongtan and Yuanmou samples to be two separate species of a single genus. The valid names for these species are Lufengpithecus lufengensis(from Shihuiba)and L. keiyuanensis(from Xiaolongtan and Yuanmou). From a phylogenetic perspective, the present available evidence suggests that lufengensis is either a primitive hominid that represents the sister taxon of the Ponginae + Homininae or a primitive sister taxon to the Ponginae. We tend to favor the second alternative, but acknowledge that a more comprehensive comparative analysis is needed to substantiate the phylogenetic and taxonomic affinities of Lufengpithcus. Importantly, the fossil apes of Yunnan provide a unique temporal perspective in the study on the evolution history of hominoeds. Their continuous occurrence during the Late Miocene and Pliocene(8Ma~2Ma), when hominoids became extinct throughout the rest of Eurasia, suggests that South China(and presumably Southeast Asia in general)was an important refugium for hominoids, including the ancestors of the orang-utans and gibbons. The uplift of the Tibetan plateau and its impact on regional climatic conditions may be an important factor in isolating the hominoids geographically and ecologically. We speculate that the changed climatic condition in the Mid-Pliocene and possibly the arrival of Homo soon after may have precipitated the regional extinction of large hominoids in South China and Southeast Asia mainland.
引文
[1]吴汝康,云南开远发现的森林古猿牙齿化石.古脊椎动物学报,1957,1:25~32.
[2]吴汝康,云南开远森林古猿新材料.古脊椎动物学报,1958,2:32~42.
[3]吴汝康,森林古猿牙齿化石在中国的首次发现及其意义,古生物学报,1958,6(2):117~121.
[4]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-颅骨的形态与比较.人类学学报,1983,2(1):1~10.
[5]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-下颌骨的形态与比较.人类学学报,1984,3(1):1~10.
[6]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-牙齿的形态与比较.人类学学报,1985,4(3):197~204.
[7]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的关系及其系统地位.人类学学报,1986,5(1):1~30.
[8]吴汝康,禄丰大猿化石分类的修订.人类学学报,1987,6(4):265~271.
[9]吴汝康,王令红,禄丰古猿的两性差别.人类学学报,1987.6(3):169~174.
[10]张兴永,云南开远新发现的腊玛古猿化石.人类学学报.1987,6(2):81~86.
[11]Kelley,J.,&Pilbeam,D .Thedryopithecines:taxonomy,anatomyphylogenyofMiocenelargehominoids.In(D.R.Swindler&J.Erwin,Eds)ComparativePrimateBiology,Volume1:Systematics,EvolutionandAnatomy,pp.361-411.NewYork:AlanR.Liss.1986.
[12]Schwartz,J.H ,Lufengpithecusandhominoidphylogeny:problemsindelineatingandevaluatingphylogeneticallyrelevantcharacters.In(D.R.Begun,C.V.Ward&M.D.Rose,Eds)Function,phylogenyandFossils:MiocensHominoidEvolutionandAdaptations,pp.363-388,NewYork:PlenumPress.1997.
[13]Ho,C.K .HumanoriginsinAsia?Hum.Evol.1988,3,357~365.
[14]Ho,C.K .AnewPliocenehominoidskullfromYunamousouthwestChina.Hum.Evol.1990,5,309~318.
[15]Wu,X.&Poirier,F.E.HumanEvolutioninChina:AMetricDescriptionofthefossilsandaReviewoftheSites.NewYork:OxfordUniversityPress.1995.
[16]董危.云南开远小龙潭动物群性质及其时代的进一步探讨.古脊椎动物学报,1987,25(2):116~123.
[17]张兴永,郑良.滇中高原与人类起源.云南社会科学1981,64(3):44~50.
[18]Qiu,Z.&Qiu,Z.ChronologicalsequenceandsubdivisionofChineseNeogenemammalianFaunas.Palaeogeogr.,Palaeoclimatol.,Palaeoecol.1995,166,41~70.
[19]Pickford,M.&Liu,L.RevisionoftheMioceneSuidaeofXiaolongtan(Kaiyuan),China.Boll.Soc.Paleontol.Ital.2001,40,275~283.
[20]尹济云.元谋盆地晚新生代地层古地磁地层年代学及其古生物意义.云南地质,1994,13(3):306~311,
[21]Simons,E.L.&Pilbeam,D.R.PreliminaryrevisionoftheDryopithecinae.Foliaprimatal.1965,3,81~152.
[22]Delson,E.Vertebratepaleontology,especiallyofnon-humanprimates,inChina.In(W.W.Howells&P.J.Tsuchitani,Eds)paleoanthropologyinthepeople'sRepublicofChina,pp.40~65.CSCPRC,ReportNo.4.Washington,D.C.:NationalAcademyofSciences.1997.
[23]徐庆华,陆庆伍.禄丰腊玛古猿下颌骨化石,科学通报,第9期,1978,554~556.
[24]张兴永,郑良,肖明华.从开远腊玛古猿的形态特征再论滇中高原与人类起源.云南社会科学,1983,66(1):83~88.
[25]吴汝康,吴新智,张森水主编,中国的远古人类.1989,245~266.科学出版社.
[26]徐庆华,陆庆五.云南禄丰发现的腊玛古猿和西瓦古猿下颌骨.古脊椎动物与古人类,1979,17(1):1~11.
[27]祁国琴.禄丰古猿化石地点化石概述.人类学学报,1985,4(1):55~69.
[28]肖明华.云南禄丰发现的古猿肩胛骨化石.云南省博物馆三十周年纪念文集,1981,41~44.
[29]林一璞等.禄丰西瓦古猿桡骨.地质论评,1987,33(1):4.
[30]Flynn,L.J.&Qi,G..AgeoftheLufeng,China,hominoidlocality.Nature1982,298,746~747.
[31]Harrison,T.&Gu,Y.TaxonomyandphylogeneticrelationshipsofearlyMiocenecatarrhinesfromSihong,China.f.hum.Evol.1999,37,225~277.
[32]刘武,郑良,姜础.元谋古猿牙齿测量数据的统计分析及其在分类研究上的意义.科学通报,1999,44(23),2481~2488.
[33]高峰.元谋古猿系统发育地位及其在人类起源研究中的意义."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集,云南科技出版社,1998,50~65.
[34]周国兴.元谋盆地人类化石及其文化遗存的研究."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集,云南科技出版社,1998,18~32.
[35]宗冠福.元谋盆地长鼻类动物化石.元谋盆地第四纪地质与古人类.北京,科学出版社.1991,178~180.
[36]吉学平.云南古猿与早期人类起源."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集.云南科技出版社,1998,44~49.
[37]潘悦容.中国新第三纪中-小型猿类化石及其意义.人类学学报,1998,17(4):283~292.
[38]潘悦容.云南元谋小河地区的灵长类化石."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集.云南科技出版社,1998,33~37.
[39]吉学平,张兴永.长鼻目.和志强主编,元谋古猿,云南科技出版社,1997,89~93.
[40]江能人,孙荣,梁其
[2]吴汝康,云南开远森林古猿新材料.古脊椎动物学报,1958,2:32~42.
[3]吴汝康,森林古猿牙齿化石在中国的首次发现及其意义,古生物学报,1958,6(2):117~121.
[4]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-颅骨的形态与比较.人类学学报,1983,2(1):1~10.
[5]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-下颌骨的形态与比较.人类学学报,1984,3(1):1~10.
[6]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的形态特征及其系统关系-牙齿的形态与比较.人类学学报,1985,4(3):197~204.
[7]吴汝康,徐庆华,陆庆伍,腊玛古猿和西瓦古猿的关系及其系统地位.人类学学报,1986,5(1):1~30.
[8]吴汝康,禄丰大猿化石分类的修订.人类学学报,1987,6(4):265~271.
[9]吴汝康,王令红,禄丰古猿的两性差别.人类学学报,1987.6(3):169~174.
[10]张兴永,云南开远新发现的腊玛古猿化石.人类学学报.1987,6(2):81~86.
[11]Kelley,J.,&Pilbeam,D .Thedryopithecines:taxonomy,anatomyphylogenyofMiocenelargehominoids.In(D.R.Swindler&J.Erwin,Eds)ComparativePrimateBiology,Volume1:Systematics,EvolutionandAnatomy,pp.361-411.NewYork:AlanR.Liss.1986.
[12]Schwartz,J.H ,Lufengpithecusandhominoidphylogeny:problemsindelineatingandevaluatingphylogeneticallyrelevantcharacters.In(D.R.Begun,C.V.Ward&M.D.Rose,Eds)Function,phylogenyandFossils:MiocensHominoidEvolutionandAdaptations,pp.363-388,NewYork:PlenumPress.1997.
[13]Ho,C.K .HumanoriginsinAsia?Hum.Evol.1988,3,357~365.
[14]Ho,C.K .AnewPliocenehominoidskullfromYunamousouthwestChina.Hum.Evol.1990,5,309~318.
[15]Wu,X.&Poirier,F.E.HumanEvolutioninChina:AMetricDescriptionofthefossilsandaReviewoftheSites.NewYork:OxfordUniversityPress.1995.
[16]董危.云南开远小龙潭动物群性质及其时代的进一步探讨.古脊椎动物学报,1987,25(2):116~123.
[17]张兴永,郑良.滇中高原与人类起源.云南社会科学1981,64(3):44~50.
[18]Qiu,Z.&Qiu,Z.ChronologicalsequenceandsubdivisionofChineseNeogenemammalianFaunas.Palaeogeogr.,Palaeoclimatol.,Palaeoecol.1995,166,41~70.
[19]Pickford,M.&Liu,L.RevisionoftheMioceneSuidaeofXiaolongtan(Kaiyuan),China.Boll.Soc.Paleontol.Ital.2001,40,275~283.
[20]尹济云.元谋盆地晚新生代地层古地磁地层年代学及其古生物意义.云南地质,1994,13(3):306~311,
[21]Simons,E.L.&Pilbeam,D.R.PreliminaryrevisionoftheDryopithecinae.Foliaprimatal.1965,3,81~152.
[22]Delson,E.Vertebratepaleontology,especiallyofnon-humanprimates,inChina.In(W.W.Howells&P.J.Tsuchitani,Eds)paleoanthropologyinthepeople'sRepublicofChina,pp.40~65.CSCPRC,ReportNo.4.Washington,D.C.:NationalAcademyofSciences.1997.
[23]徐庆华,陆庆伍.禄丰腊玛古猿下颌骨化石,科学通报,第9期,1978,554~556.
[24]张兴永,郑良,肖明华.从开远腊玛古猿的形态特征再论滇中高原与人类起源.云南社会科学,1983,66(1):83~88.
[25]吴汝康,吴新智,张森水主编,中国的远古人类.1989,245~266.科学出版社.
[26]徐庆华,陆庆五.云南禄丰发现的腊玛古猿和西瓦古猿下颌骨.古脊椎动物与古人类,1979,17(1):1~11.
[27]祁国琴.禄丰古猿化石地点化石概述.人类学学报,1985,4(1):55~69.
[28]肖明华.云南禄丰发现的古猿肩胛骨化石.云南省博物馆三十周年纪念文集,1981,41~44.
[29]林一璞等.禄丰西瓦古猿桡骨.地质论评,1987,33(1):4.
[30]Flynn,L.J.&Qi,G..AgeoftheLufeng,China,hominoidlocality.Nature1982,298,746~747.
[31]Harrison,T.&Gu,Y.TaxonomyandphylogeneticrelationshipsofearlyMiocenecatarrhinesfromSihong,China.f.hum.Evol.1999,37,225~277.
[32]刘武,郑良,姜础.元谋古猿牙齿测量数据的统计分析及其在分类研究上的意义.科学通报,1999,44(23),2481~2488.
[33]高峰.元谋古猿系统发育地位及其在人类起源研究中的意义."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集,云南科技出版社,1998,50~65.
[34]周国兴.元谋盆地人类化石及其文化遗存的研究."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集,云南科技出版社,1998,18~32.
[35]宗冠福.元谋盆地长鼻类动物化石.元谋盆地第四纪地质与古人类.北京,科学出版社.1991,178~180.
[36]吉学平.云南古猿与早期人类起源."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集.云南科技出版社,1998,44~49.
[37]潘悦容.中国新第三纪中-小型猿类化石及其意义.人类学学报,1998,17(4):283~292.
[38]潘悦容.云南元谋小河地区的灵长类化石."元谋人"发现三十周年纪念暨古人类国际学术研讨会文集.云南科技出版社,1998,33~37.
[39]吉学平,张兴永.长鼻目.和志强主编,元谋古猿,云南科技出版社,1997,89~93.
[40]江能人,孙荣,梁其
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