高原鼢鼠(Eospalax baileyi)头骨形态的地理分化
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摘要
运用单因素方差分析、主成分分析、判别分析和聚类分析,对高原鼢鼠(Eospalax baileyi)4个地区(临潭、天祝、玛曲和碌曲)8个地理群体的头骨形态特征进行综合分析,比较13个特征变量的地理分异,挖掘形态特征的主要变量,并探讨其形态变异与环境的关系。结果发现,群体间13个特征变量均存在显著差异,主成分分析表明13个头骨特征变量可用3个主成分来表述,其累计贡献率达96.981%。颅全长、上齿列长及齿隙长是高原鼢鼠头骨发生分化的主要变量。天祝种群与碌曲种群,及玛曲种群与临潭种群都有重叠,其中碌曲的两个种群相似性最高,玛曲种群和临潭种群与其他种群几乎无重叠,显示完全分化,聚类分析结果与判别分析结果一致。头骨形态与地理因子的相关性分析发现头骨形态大小与海拔成显著的正相关关系。综合分析认为地理隔离、栖息地海拔因子影响下的生态条件等是高原鼢鼠头骨分化的主要因素。
Morphological variation stems from a number of sources,including environmental factors. The skull is the most complex part of animal skeleton. Studies on skull morphological variations can help uncover the taxonomic status of a species,age and other ecological characteristics of the populations. Qinghai-Tibetan Plateau( QTP) is a region with high degree of environmental heterogeneity and harbors a rich diversity of indigenous fauna. Plateau zokor belong to the genus Eospalax and are endemic to QTP. Zokors have a long evolutionary history that has resulted in specialized characteristics such as reduced eye size and sight,strong adaptation to hypoxia,and foreleg structure,and strong claws for digging. These make this species an ideal model for testing the role of environmental factors in shaping morphological features. The skulls of plateau zokor from eight geographical populations were collected from the northeast of QTP in Lintan,Tianzhu,Maqu,and Luqu of the Gansu Province. The morphometric measurements were taken and the geographic division of 13 morphological characteristics was compared and analyzed by a one-way analysis of variance and discriminant function analysis. Theenvironmental influences on morphological differentiationwere discussed. The results indicated that there were significant differences in four morphological characteristics and highly significant differences in two morphological characteristics between the Lintan and Luqu groups,but there was only one significant morphological characteristic between the Maqu and Lintan groups,Lintan and Tianzhu groups,and Maqu and Luqu groups. Principal components analyses showed that the 13 main traits were represented by three principal components,and contributed to 96.981% of variations cumulatively stepwise discriminant function analysis showed the diastema length and total length had the greatest contribution to skull shape variation. A significant positive correlation between zokor skull size and altitude was detected. These findings suggested that geographical isolation and ecological conditions under the influence of altitude was the main factor shaping zokor skull differentiation.
Morphological variation stems from a number of sources,including environmental factors. The skull is the most complex part of animal skeleton. Studies on skull morphological variations can help uncover the taxonomic status of a species,age and other ecological characteristics of the populations. Qinghai-Tibetan Plateau( QTP) is a region with high degree of environmental heterogeneity and harbors a rich diversity of indigenous fauna. Plateau zokor belong to the genus Eospalax and are endemic to QTP. Zokors have a long evolutionary history that has resulted in specialized characteristics such as reduced eye size and sight,strong adaptation to hypoxia,and foreleg structure,and strong claws for digging. These make this species an ideal model for testing the role of environmental factors in shaping morphological features. The skulls of plateau zokor from eight geographical populations were collected from the northeast of QTP in Lintan,Tianzhu,Maqu,and Luqu of the Gansu Province. The morphometric measurements were taken and the geographic division of 13 morphological characteristics was compared and analyzed by a one-way analysis of variance and discriminant function analysis. Theenvironmental influences on morphological differentiationwere discussed. The results indicated that there were significant differences in four morphological characteristics and highly significant differences in two morphological characteristics between the Lintan and Luqu groups,but there was only one significant morphological characteristic between the Maqu and Lintan groups,Lintan and Tianzhu groups,and Maqu and Luqu groups. Principal components analyses showed that the 13 main traits were represented by three principal components,and contributed to 96.981% of variations cumulatively stepwise discriminant function analysis showed the diastema length and total length had the greatest contribution to skull shape variation. A significant positive correlation between zokor skull size and altitude was detected. These findings suggested that geographical isolation and ecological conditions under the influence of altitude was the main factor shaping zokor skull differentiation.
引文
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[12]李松,杨君兴,蒋学龙,王应祥.中国巨松鼠Ratufa bicolor(Sciuridae:Ratufinae)头骨形态的地理学变异.兽类学报,2008,28(2):201-206.
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[17]陶燕铎,樊乃昌,景增春.高原鼢鼠对草场的危害及防治阈值的探讨.中国媒介生物学及控制杂志,1990,1(2):103-106.
[18]周立志,李迪强,王秀磊,马强.三江源自然保护区鼠害类型、现状和防治策略.安徽大学学报:自然科学版,2002,26(2):87-96.
[19]苏军虎,刘荣堂,纪维红,焦婷,蔡卓山,花立民.我国草地鼠害防治与研究的发展阶段及特征.草业科学,2013,30(7):1116-1123.
[20]苏军虎,Ji W H,南志标,刘荣堂,王静,徐长林.鼢鼠亚科Mysopalacinae动物系统学研究现状与展望.动物学杂志,2015,50(4):649-658.
[21]唐利洲,张同作,苏建平.高原鼢鼠(Myospalax baileyi)不同地理种群的形态变异.兽类学报,2009,29(2):178-184.
[22]鲁庆彬,张阳,周材权.甘肃鼢鼠不同地理种群的形态变异分析.兽类学报,2013,33(2):193-199.
[23]杨奇森,夏霖,马勇,冯祚建,全国强.兽类头骨测量标准I:基本量度.动物学杂志,2005,40(3):50-56.
[24]薛薇.统计分析与SPSS的应用(第二版).北京:中国人民大学出版社,2008:136-312.
[25]Geng Y,Wang Z H,Liang C Z,Fang J Y,Baumann F,Kühn P,Scholten T,He J S.Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands.Global Ecology and Biogeography,2012,21(4):416-427.
[26]Wright S.Isolation by distance.Genetics,1943,28(2):114-138.
[27]Trizio I,Crestanello B,Galbusera P,Wauters L A,Tosi G,Matthysen E,Hauffe H C.Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels(Sciurus vulgaris)in the Italian Alps.Molecular Ecology,2005,14(2):469-481.
[28]蔡振媛,张同作,慈海鑫,唐利洲,连新明,刘建全,苏建平.高原鼢鼠线粒体谱系地理学和遗传多样性.兽类学报,2007,27(2):130-137.
[29]李惠萍,张放,韩崇选.陕西草兔头骨形态的地理学分化.西北林学院学报,2011,26(1):109-112,127-127.
[30]Therry L,Gyulavári H A,Schillewaert S,Bonte D,Stoks R.Integrating large-scale geographic patterns in flight morphology,flight characteristics and sexual selection in a range-expanding damselfly.Ecography,2014,37(10):1012-1021.
[31]何锴,白明,万韬,李权,王应祥,蒋学龙.白尾鼹(鼹科:哺乳纲)下颌骨几何形态测量分析及地理分化研究.兽类学报,2013,33(1):7-17.
[32]Borges L R,Maestri R,Kubiak B B,Galiano D,Fornel R,Freitas T R O.The role of soil features in shaping the bite force and related skull and mandible morphology in the subterranean rodents of genus Ctenomys(Hystricognathi:Ctenomyidae).Journal of Zoology,2017,301(2):108-117.
[2]Dresssino V,Pucciarelli H M.Cranial growth in Saimiri sciureus(Cebidae)and its alteration by nutritional factors:a longitudinal study.American Journal of Physical Anthropology,1997,102(4):545-554.
[3]Stumpp R,Fuzessy L,Paglia A P.Environment drivers acting on rodent rapid morphological change.Journal of Mammalian Evolution,2016,doi:10.1007/s10914-016-9369-2.
[4]Marcy A E,Hadly E A,Sherratt E,Garland K,Weisbecker V.Getting a head in hard soils:convergent skull evolution and divergent allometric patterns explain shape variation in a highly diverse genus of pocket gophers(Thomomys).BMC Evolutionary Biology,2016,16:207.
[5]Assis A P A,Rossoni D M,Patton J L,Marroig G.Evolutionary processes and its environmental correlates in the cranial morphology of western chipmunks(Tamias).Evolution,2016,doi:10.1111/evo.13137.
[6]邰发道,孙儒泳,王廷正,周宏伟.五种鼠的脑和头骨形态及其生态关系.动物学研究,2001,22(6):472-477.
[7]熊文华,张知彬.饶阳地区三种农田啮齿类头骨形态比较及性二型.兽类学报,2007,27(3):280-283.
[8]Haas A,Richards S J.Correlations of cranial morphology,ecology,and evolution in Australian suctorial tadpoles of the genera Litoria and Nyctimystes(Amphibia:Anura:Hylidae:Pelodryadinae).Journal of Morphology,1998,238(2):109-141.
[9]Silva J M D,Carne L,John Measey G,Herrel A,Tolley K A.The relationship between cranial morphology,bite performance,diet and habitat in a radiation of dwarf chameleon(Bradypodion).Biological Journal of the Linnean Society,2016,119(1):52-67.
[10]葛德燕,吕雪霏,夏霖,黄乘明,杨奇森.草兔胚后发育过程中头骨大小与形状变化的几何形态学分析(兔形目:兔科).兽类学报,2012,32(1):12-24.
[11]高文荣,王政昆,姜文秀,朱万龙.云南高山姬鼠头骨的几何形态学研究.生态学报,2016,36(6):1756-1764.
[12]李松,杨君兴,蒋学龙,王应祥.中国巨松鼠Ratufa bicolor(Sciuridae:Ratufinae)头骨形态的地理学变异.兽类学报,2008,28(2):201-206.
[13]阿尔孜古力·沙塔尔,买热帕提·帕拉提,迪力夏提·阿不力孜,马合木提·哈力克.塔里木兔不同地理种群头骨形态特征的比较研究.新疆农业科学,2010,47(8):1627-1631.
[14]张明春,张泽钧,母华强,黄小富,胡锦矗.川西四大山系间高山姬鼠头骨特征的地理分化.兽类学报,2008,28(4):395-401.
[15]Norris R W,Zhou K Y,Zhou C Q,Yang G,William Kilpatrick C,Honeycutt R L.The phylogenetic position of the zokors(Myospalacinae)and comments on the families of muroids(Rodentia).Molecular Phylogenetics and Evolution,2004,31(3):972-978.
[16]Zhang Y M,Zhang Z B,Liu J K.Burrowing rodents as ecosystem engineers:the ecology and management of plateau zokors Myospalax fontanierii in alpine meadow ecosystems on the Tibetan Plateau.Mammal Review,2003,33(3/4):284-294.
[17]陶燕铎,樊乃昌,景增春.高原鼢鼠对草场的危害及防治阈值的探讨.中国媒介生物学及控制杂志,1990,1(2):103-106.
[18]周立志,李迪强,王秀磊,马强.三江源自然保护区鼠害类型、现状和防治策略.安徽大学学报:自然科学版,2002,26(2):87-96.
[19]苏军虎,刘荣堂,纪维红,焦婷,蔡卓山,花立民.我国草地鼠害防治与研究的发展阶段及特征.草业科学,2013,30(7):1116-1123.
[20]苏军虎,Ji W H,南志标,刘荣堂,王静,徐长林.鼢鼠亚科Mysopalacinae动物系统学研究现状与展望.动物学杂志,2015,50(4):649-658.
[21]唐利洲,张同作,苏建平.高原鼢鼠(Myospalax baileyi)不同地理种群的形态变异.兽类学报,2009,29(2):178-184.
[22]鲁庆彬,张阳,周材权.甘肃鼢鼠不同地理种群的形态变异分析.兽类学报,2013,33(2):193-199.
[23]杨奇森,夏霖,马勇,冯祚建,全国强.兽类头骨测量标准I:基本量度.动物学杂志,2005,40(3):50-56.
[24]薛薇.统计分析与SPSS的应用(第二版).北京:中国人民大学出版社,2008:136-312.
[25]Geng Y,Wang Z H,Liang C Z,Fang J Y,Baumann F,Kühn P,Scholten T,He J S.Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands.Global Ecology and Biogeography,2012,21(4):416-427.
[26]Wright S.Isolation by distance.Genetics,1943,28(2):114-138.
[27]Trizio I,Crestanello B,Galbusera P,Wauters L A,Tosi G,Matthysen E,Hauffe H C.Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels(Sciurus vulgaris)in the Italian Alps.Molecular Ecology,2005,14(2):469-481.
[28]蔡振媛,张同作,慈海鑫,唐利洲,连新明,刘建全,苏建平.高原鼢鼠线粒体谱系地理学和遗传多样性.兽类学报,2007,27(2):130-137.
[29]李惠萍,张放,韩崇选.陕西草兔头骨形态的地理学分化.西北林学院学报,2011,26(1):109-112,127-127.
[30]Therry L,Gyulavári H A,Schillewaert S,Bonte D,Stoks R.Integrating large-scale geographic patterns in flight morphology,flight characteristics and sexual selection in a range-expanding damselfly.Ecography,2014,37(10):1012-1021.
[31]何锴,白明,万韬,李权,王应祥,蒋学龙.白尾鼹(鼹科:哺乳纲)下颌骨几何形态测量分析及地理分化研究.兽类学报,2013,33(1):7-17.
[32]Borges L R,Maestri R,Kubiak B B,Galiano D,Fornel R,Freitas T R O.The role of soil features in shaping the bite force and related skull and mandible morphology in the subterranean rodents of genus Ctenomys(Hystricognathi:Ctenomyidae).Journal of Zoology,2017,301(2):108-117.