收稿日期: 2020-03-24
修回日期: 2020-05-20
网络出版日期: 2021-03-09
基金资助
国家自然科学基金项目(41877455);国家自然科学基金项目(41761018);青海省自然科学基金青年项目(2017-ZJ-931Q);中国科学院战略性先导科技专项“泛第三极环境变化与绿色丝绸之路建设”(XDA2004010101)
Characteristics and environmental implications of dung pollen assemblages of grazing livestock in alpine meadow in the eastern Qinghai-Tibetan Plateau
Received date: 2020-03-24
Revised date: 2020-05-20
Online published: 2021-03-09
青藏高原是我国重要的高寒草甸分布区和畜牧区,畜牧活动对区域植被和生态的影响作用受到众多学者的广泛关注。通过采集青藏高原东部高寒草甸区共49个放牧家畜粪样品（牦牛粪样品30个、马粪样品11个、羊粪样品8个）,并在区域植被调查的基础上,对粪样品开展了花粉分析。结果表明：这3种类型的粪样品花粉组合以莎草科（Cyperaceae）、禾本科（Poaceae）、龙胆科（Gentianaceae）、毛茛科（Ranunculaceae）花粉为主,可较好地反映区域植被群落特征,食草动物粪花粉中有少量谷物类禾本科花粉出现反映了当地居民少量耕种燕麦用来饲养牲畜。3种放牧家畜在采食习性上存在一定差异,牦牛采食的植物种类比羊和马略多,牦牛和马对杂草类禾本科和唐松草属植物的采食率低于羊,对龙胆科、菊科和蒿属植物的采食率明显高于羊。放牧家畜通过选择性采食适口性好的莎草科和禾本科优势植物种类直接影响高寒草甸区植被群落结构和功能。研究区粪花粉组合与表土花粉组合对比发现,乔木花粉和紫菀属花粉含量在表土样品中显著高于粪样品,而禾本科和龙胆科花粉含量在粪样品中高于表土样品,上述差异指示粪花粉更能反映区域植被群落特征,放牧家畜选择性采食导致粪花粉与表土花粉组合有所区别。本研究为评估青藏高原放牧家畜采食习性以及畜牧活动对区域植被的影响提供了新的研究思路和手段,也为今后在青藏高原开展考古遗址中保存的粪样品花粉分析,据此重建古植被群落和古人类生产活动信息,提供了重要的现代过程依据和借鉴资料。
段荣蕾,侯光良,魏海成,许清海,高靖易 . 青藏高原东部高寒草甸区放牧家畜粪花粉组合特征及其环境指示意义[J]. 干旱区地理, 2021 , 44(1) : 229 -239 . DOI: 10.12118/j.issn.1000–6060.2021.01.24
The Qinghai-Tibetan Plateau is an important area of alpine meadow pasture and livestock husbandry in China. The impact of grazing activity on the vegetation and ecology in the Qinghai-Tibetan Plateau has attracted widespread attention from many scholars. To study the impact of grazing activity on regional vegetation, we collected 49 dung samples of grazing livestock (comprising 30 yak dung, 11 horse dung, and 8 sheep dung) from the typical alpine meadow in the eastern Qinghai-Tibetan Plateau. Our study area extended from 34°30′08″- 35°06′28″N, 101°30′57″-101°42′04″E, with elevations ranging between 3456 m and 3815 m. The vegetation communities in the sampling sites were also investigated during the field work. Principal component analysis was used to explore the characteristics of the different dung samples and pollen taxa. Meanwhile, the pollen assemblage of the dung samples was compared with the topsoil pollen assemblages in the study area to examine the selective grazing characteristics of the different livestock in the Qinghai-Tibetan Plateau. These results indicate that the pollen assemblages in the three types of livestock’s dung samples were dominated by Cyperaceae, Poaceae, Gentianaceae, and Ranunculaceae, generally reflecting the characteristics of the regional plant communities. However, there are some differences in the feeding habits of the three kinds of grazing livestock: the feeding rates of yaks and horses to Poaceae and Thalictrum species are lower than those of sheep, and those to Gentianaceae, Compositae, and Artemisia species are significantly higher than those of sheep. In addition, cereal Poaceae pollen appeared in the dung samples, reflecting the behavior of the local residents in cultivating oats to raise their livestock. Grazing livestock directly affects the vegetation community structure of alpine meadow through selective feeding of the dominant plant species of Cyperaceae and Poaceae. The comparison of the pollen assemblages of the dung and topsoil samples in the study area indicates that the contents of Cyperaceae, Polygonaceae, Ranunculaceae, Taraxacum, and Artemisia were similar. However, the contents of Poaceae and Gentianaceae in the dung samples were higher than those in the topsoil samples, and the content of Aster in the topsoil samples was higher than that in dung samples, suggesting the selective feeding characteristics of the grazing livestock. Additionally, the content of arboreal pollen, including Betula, Pinus, Picea, and Hippophae, in the topsoil samples was higher than those in the dung samples, indicating that the dung pollen assemblages could better reflect the characteristics of the local vegetation community. This study provides a new research approach for assessing the grazing habits of livestock and impact of grazing activity on regional vegetation in the Qinghai-Tibetan Plateau. Additionally, the study of pollen assemblages in modern dung samples provides an important reference for the fossil dung pollen assemblages in the archaeological sites to infer the past vegetation and ancient humans’ subsistence strategy on the Qinghai-Tibetan Plateau.
| [1] | 姚檀栋, 陈发虎, 崔鹏, 等. 从青藏高原到第三极和泛第三极[J]. 中国科学院院刊, 2017,32(9):924-931. |
| [1] | [ Yao Tandong, Chen Fahu, Cui Peng, et al. From Tibetan Plateau to Third Pole and Pan-Third Pole[J]. Bulletin of Chinese Academy of Sciences, 2017,32(9):924-931. ] |
| [2] | 孙鸿烈, 郑度, 姚檀栋, 等. 青藏高原国家生态安全屏障保护与建设[J]. 地理学报, 2012,67(1):3-12. |
| [2] | [ Sun Honglie, Zheng Du, Yao Tandong, et al. Protection and construction of the national ecological security shelter zone on Tibetan Plateau[J]. Acta Geographica Sinica, 2012,67(1):3-12. ] |
| [3] | Yao T D, Thompson L G, Mosbrugger V, et al. Third pole environment (TPE)[J]. Environmental Development, 2012(3):52-64. |
| [4] | Schwalb A, Schütt B, Fang X. Climate evolution and environmental response on the Tibetan Plateau[J]. Quaternary International, 2010(218):1-2. |
| [5] | Qiu J. China: The third pole[J]. Nature News, 2008,454(7203):393-396. |
| [6] | 谢高地, 鲁春霞, 肖玉, 等. 青藏高原高寒草地生态系统服务价值评估[J]. 山地学报, 2003(1):50-55. |
| [6] | [ Xie Gaodi, Lu Chunxia, Xiao Yu, et al. The economic evaluation of grassland ecosystem services in Qinghai-Tibet Plateau[J]. Mountain Research, 2003(1):50-55. ] |
| [7] | 车宗玺, 闫明, 何同弟, 等. 祁连山西段草地土壤氮矿化及影响因子的关系研究[J]. 干旱区地理, 2018,41(1):92-98. |
| [7] | [ Che Zongxi, Yan Ming, He Tongdi, et al. Soil nitrogen mineralization and its influence factors in grassland in western Qilian Mountains[J]. Arid Land Geography, 2018,41(1):92-98. ] |
| [8] | Li G, Zhang Z, Shi L L, et al. Effects of different grazing intensities on soil C, N, and P in an alpine meadow on the Qinghai-Tibetan Plateau, China[J]. International Journal of Environmental Research and Public Health, 2018,15(11):2584, doi: 10.3390/ijerph15112584 |
| [9] | 杨晶晶, 吐尔逊娜依·热依木, 张青青, 等. 放牧强度对天山北坡中段山地草甸植被群落特征的影响[J]. 草业科学, 2019,36(8):1953-1961. |
| [9] | [ Yang Jingjing, Tuerxunnayi Reyimu, Zhang Qingqing, et al. Effects of grazing intensity on plant community characteristics in mountain meadows in the middle section of the northern slope of the Tianshan Mountains[J]. Pratacultural Science, 2019,36(8):1953-1961. ] |
| [10] | Gao Y, Luo P, Wu N, et al. Biomass and nitrogen responses to grazing intensity in an alpine meadow on the eastern Tibetan Plateau[J]. Polish Journal of Ecology, 2007,55(3):469. |
| [11] | 乌云娜, 张凤杰, 盐见正卫. 放牧梯度上群落物种数与物种组成结构异质性的数量分析[J]. 干旱区地理, 2011,34(6):904-911. |
| [11] | [ Wu Yunna, Zhang Fengjie, Shiyomi Masae. Quantitative analysis of spatial heterogeneity in species richness and composition in grassland vegetation with different grazing intensities in the northeast Inner Mongolian steppe[J]. Arid Land Geography, 2011,34(6):904-911. ] |
| [12] | 刘伟, 王溪, 刘季科, 等. 家畜实验放牧强度对啮齿动物群落多样性作用的研究[J]. 青海畜牧兽医杂志, 1990(5):3-6. |
| [12] | [ Liu Wei, Wang Xi, Liu Jike, et al. Effects of experimental grazing intensity on rodent community diversity[J]. Chinese Qinghai Journal of Animal and Veterinary Sciences, 1990(5):3-6. ] |
| [13] | 周立华, 侯彩霞. 北方农牧交错区草原利用与禁牧政策的关键问题研究[J]. 干旱区地理, 2019,42(2):354-362. |
| [13] | [ Zhuo Lihua, Hou Caixia. Key problems of grassland utilization and the graze ban policy in farming-pastoral ecotone of northern China[J]. Arid Land Geography, 2019,42(2):354-362. |
| [14] | 秦景秀, 郝兴明, 张颖, 等. 气候变化和人类活动对干旱区植被生产力的影响[J]. 干旱区地理, 2020,43(1):117-125. |
| [14] | [ Qin Jingxiu, Hao Xingming, Zhang Ying, et al. Effects of climate change and human activities on vegetation productivity in arid areas[J]. Arid Land Geography, 2020,43(1):117-125. ] |
| [15] | Diaz S, Lavorel S, Mcintyre S, et al. Plant trait responses to grazing: A global synjournal[J]. Global Change Biology, 2007,13(2):313-341. |
| [16] | Marcelo S, Mario G, Avi P, et al. Vegetation response to grazing management in a Mediterranean herbaceous community: A functional group approach[J]. Journal of Applied Ecology, 2000,37(2):224-237. |
| [17] | 李金花, 李镇清, 任继周. 放牧对草原植物的影响[J]. 草业学报, 2002,11(1):4-11. |
| [17] | [ Li Jinhua, Li Zhenqing, Ren Jizhou. The effects of grazing on grassland plants[J]. Acta Prataculturae Sinica, 2002,11(1):4-11. ] |
| [18] | 杜岩功, 梁东营, 曹广民, 等. 放牧强度对嵩草草甸草毡表层及草地营养和水分利用的影响[J]. 草业学报, 2008,17(3):146-150. |
| [18] | [ Du Yangong, Liang Dongying, Cao Guangmin, et al. Effect on incrassation of mattic epipedon and pasture nutrient-moisture using by grazing in alpine Kobresia meadow[J]. Acta Prataculturae Sinica, 2008,17(3):146-150. ] |
| [19] | 李永宏, 汪诗平. 内蒙古细毛羊日食量及对典型草原牧草的食性测定[J]. 草业学报, 1998,7(1):50-53. |
| [19] | [ Li Yonghong, Wang Shiping. Grazing intake of inner Mongolian fine-wool sheep and their preference for grassland forage species[J]. Acta Prataculturae Sinica, 1998,7(1):50-53. ] |
| [20] | 汪诗平. 不同放牧率下绵羊的食性及食物多样性与草地植物多样性间的关系[J]. 生态学报, 2001,21(2):237-243. |
| [20] | [ Wang Shiping. The dietary composition of fine wool sheep under different stocking rates and relationship between dietary diversity and plant diversity in Inner Mongolia steppe[J]. Acta Ecologica Sinica, 2001,21(2):237-243. ] |
| [21] | 王旭, 王德利, 刘颖, 等. 不同放牧率下绵羊的采食量与食性选择研究[J]. 东北师大学报(自然科学版), 2002,34(1):36-40. |
| [21] | [ Wang Xu, Wang Deli, Liu Ying, et al. The study on the intake and dietary composition of sheep under different stocking rates[J]. Journal of Northeast Normal University, 2002,34(1):36-40. ] |
| [22] | 宋仁德, 长谷川信美, 李国梅, 等. 天然草地放牧牦牛采食行为及食性选择的研究[J]. 家畜生态学报, 2008,29(5):31-35. |
| [22] | [ Song Rende, Hasagawa Nobumi, Li Guomei, et al. Botanical composition and grazing behaviour of Qinghai yaks of plateau type in the natural rangeland[J]. Journal of Domestic Animal Ecology, 2008,29(5):31-35. ] |
| [23] | Kamler J, Homolka M. Needles in faeces: An index of quality of wild ungulate winter diet[J]. Folia Zoologica, 2011,60(1):63-69. |
| [24] | Marti C. Accuracy of fecal analysis for identifying foods of black grouse[J]. The Journal of Wildlife Management, 1982,46(3):773-777. |
| [25] | Norbury G. Microscopic analysis of herbivore diets: A problem and a solution[J]. Wildlife Research, 1988,15(1):51-57. |
| [26] | 高中信, 陈化鹏, 王筱平. 粪便分析法测定植食动物食性的评价[J]. 兽类学报, 1991,11(3):186-193. |
| [26] | [ Gao Zhongxin, Chen Huapeng, Wang Xiaoping. Evaluation of fecal analysis for determining food habits of herbivores[J]. Acta Theriologica Sinica, 1991,11(3):186-193. ] |
| [27] | 郑荣泉, 鲍毅新. 有蹄类食性研究方法及研究进展[J]. 生态学报, 2004,24(7):1532-1539. |
| [27] | [ Zheng Rongquan, Bao Yixin. Study methods and procedures for ungulate food habits[J]. Acta Ecologica Sinica, 2004,24(7):1532-1539. ] |
| [28] | 单继红, 吴建平. 食草动物食性研究的主要方法及其评价[J]. 野生动物杂志, 2005,26(3):47-49. |
| [28] | [ Shan Jihong, Wu Jianping. The review and evaluation on the study methods for herbivore food habit[J]. Chinese Journal of Wildlife, 2005,26(3):47-49. ] |
| [29] | 王开发, 王宪曾. 孢粉学概论[M]. 北京: 北京大学出版社, 1983: 1-205. |
| [29] | [ Wang Kaifa, Wang Xianzeng. Introduction to spore-pollen study[M]. Beijing: Peking University Press, 1993: 1-205. ] |
| [30] | Carrion J S. A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain[J]. Review of Palaeobotany and Palynology, 2002,120(3):217-232. |
| [31] | Basumatary S K, Mcdonald H G, Gogoi R, et al. Pollen and non-pollen palynomorph preservation in the dung of the greater one-horned rhino (Rhinoceros unicornis), and its implication to palaeoecology and palaeodietary analysis: A case study from India[J]. Review of Palaeobotany and Palynology, 2017,244:153-162. |
| [32] | 郝瑞辉, 萧家仪, 房迎三, 等. 南京汤山驼子洞鬣狗粪化石的孢粉分析[J]. 古生物学报, 2008,47(1):123-128. |
| [32] | [ He Ruihui, Xiao Jiayi, Fang Yinsan, et al. Pollen analysis of Hyaena coprolites from the Tuozi Cave at Tangshan Nanjing[J]. Acta Palaeontologica Sinica, 2008,47(1):123-128. ] |
| [33] | Thompson R S, Van Devender T R, Martin P S, et al. Shasta ground sloth (Nothrotheriops shastense Hoffstetter) at shelter cave, New Mexico: Environment, diet, and extinction[J]. Quaternary Research, 1980,14(3):360-376. |
| [34] | Alcover J A, Perez-Obiol R, Yll E I, et al. The diet of Myotragus balearicus Bate 1909 (Artiodactyla: Caprinae), an extinct bovid from the Balearic Islands: Evidence from coprolites[J]. Biological Journal of the Linnean Society, 1999,66(1):57-74. |
| [35] | Welker F, Duijm E, Der Gaag K J, et al. Analysis of coprolites from the extinct mountain goat Myotragus balearicus[J]. Quaternary Research, 2014,81(1):106-116. |
| [36] | Yll R, Carrion J S, Marra A C, et al. Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy)[J]. Palaeogeography, Palaeoclimatology, Palaeoecology, 2006,237(1):32-39. |
| [37] | 杜乃秋, 于浅黎. 周口店鬣狗(Hyaena)粪化石的孢粉分析[J]. 古脊椎动物学报, 1980,18(3):83. |
| [37] | [ Du Naiqiu, Yu Qianli. Sporopoly analysis of coprolites of Hyaena in Zhoukoudian[J]. Vertebrata Palasiatica, 1980,18(3):83. ] |
| [38] | 王文娟, 吴妍, 宋国定, 等. 灵井许昌人遗址鬣狗粪化石的孢粉和真菌孢子研究[J]. 科学通报, 2013,58(增刊1):51-56. |
| [38] | [ Wang Wenjuan, Wu Yan, Song Guoding, et al. ollen and fungi spore analysis on Hyaenid coprolite from the Xuchang Man Site, central China[J]. Chinese Science Bulletin, 2014, 58(Suppl. 1): 51-56. |
| [39] | Marcolino C P, Isaias R M, Cozzuol M A, et al. Diet of Palaeolama major (Camelidae) of Bahia, Brazil, inferred from coprolites[J]. Quaternary International, 2012,278:81-86. |
| [40] | Velázquez N J, Burry L S, Mancini M V, et al. Coprolitos de camélidos del Holoceno como indicadores paleoambientales[J]. Magallania (Punta Arenas), 2010,38(2):213-229. |
| [41] | Qiu Z, Yang Y, Shang X, et al. Paleo-environment and paleo-diet inferred from Early Bronze Age cow dung at Xiaohe Cemetery, Xinjiang, NW China[J]. Quaternary International, 2014,349:167-177. |
| [42] | Zhang Y, Van Geel B, Gosling W D, et al. Local vegetation patterns of a Neolithic environment at the site of Tianluoshan, China, based on coprolite analysis[J]. Review of Palaeobotany and Palynology, 2019,271, doi: 10.1016/j.revpalbo.2019.104101 |
| [43] | 吴慧贤, 徐海, 蓝江湖, 等. 北疆表土花粉组合及其与植被和气候的关系[J]. 生态学杂志, 2018,37(12):3499-3507. |
| [43] | [ Wu Huixian, Xu Hai, Lan Jianghu, et al. Pollen assemblages in surface soil and their relationships with vegetation and climate in northern Xinjiang[J]. Chinese Journal of Ecology, 2018,37(12):3499-3507. ] |
| [44] | 周华坤, 王启基, 赵亮, 等. 青海省泽库县草地现状与畜牧业可持续发展对策[J]. 草业科学, 2007,24(3):87-93. |
| [44] | [ Zhou Huakun, Wang Qiji, Zhao Liang, et al. The present situation of grassland and the strategies of sustainable development of animal husbandry in Zeku County, Qinghai Province[J]. Acta Prataculturae Sinica, 2007,24(3):87-93. ] |
| [45] | 徐开宇, 纳得秀, 侯琳. 黄南地区降水分布特征及人工增雨的最佳气象条件分析[J]. 青海气象, 2009(2):25-28. |
| [45] | [ Xu Kaiyu, Na Dexiu, Hou Lin. The characteristics of precipitation distribution and the optimum meteorological conditions of artificial precipitation in Huangnan area[J]. Journal of Qinghai Meteorology, 2009(2):25-28. ] |
| [46] | 杜加强, 舒俭民, 张林波. 基于NPP的黄南州自然植被对气候变化的响应[J]. 生态学杂志, 2010,29(6):1094-1102. |
| [46] | [ Du Jiaqiang, Shu Jianmin, Zhang Linbo. Responses of natural vegetation in Huangnan Prefecture of Qinghai to climate change: A study based on NPP[J]. Chinese Journal of Ecology, 2010,29(6):1094-1102. ] |
| [47] | 张国胜, 马元仓, 董立新, 等. 青海省柴达木盆地若干风速特征分析[J]. 青海气象, 2002(1):27-31. |
| [47] | [ Zhang Guosheng, Ma Yuancang, Dong Lixin, et al. Analysis of wind speed characteristics in Qaidam Basin, Qinghai Province[J]. Journal of Qinghai Meteorology, 2002(1):27-31. ] |
| [48] | 周兴明. 青海植被[M]. 西宁: 青海人民出版社, 1986: 1-199. |
| [48] | [ Zhou Xingmin. Vegetation of Qinghai Province[M]. Xining: Qinghai People’s Press, 1986: 1-199. ] |
| [49] | 侯学煜. 中国植被图集(1:1000000)[M]. 北京: 科学出版社, 2001: 1-260. |
| [49] | [ Hou Xueyu. Vegetation atlas of China: (1:1000000)[M]. Beijing: Science Press, 2001: 1-260. ] |
| [50] | 王涛, 许清海, 卢静瑶, 等. 青藏高原东北部高寒草甸夏季开花植物花粉形态研究[J]. 古生物学报, 2018,57(4):524-542. |
| [50] | [ Wang Tao, Xu Qinghai, Lu Jingyao, et al. Pollen morphol of summer-plowering species of alpine meadows in northeastern Qinghai Tibetan Plateau[J]. Acta Palaeontologica Sinica, 2018,57(4):524-542. ] |
| [51] | 王伏雄, 钱南芬, 张玉龙, 等. 中国植物花粉形态[M]. 2版. 北京: 科学出版社, 1997: 1-461. |
| [51] | [ Wang Fuxiong, Qian Nanfen, Zhang Yulong, et al. Pollen flora of China[M]. 2nded. Beijing: Science Press, 1997: 1-461. ] |
| [52] | 唐领余, 毛礼米, 舒军武, 等. 中国第四纪孢粉图鉴[M]. 北京: 科学出版社, 2016: 1-621. |
| [52] | [ Tang Lingyu, Mao Limi, Shu Junwu, et al. An illustrated handbook of Quaternary pollen and spores in China[M]. Beijing: Science Press, 2016: 1-621. ] |
| [53] | Grimm E C. TILIA software version 1.7.16[M]. Springfield USA: Illinois State Museum, Research and Collection Center, 2011. |
| [54] | Herzschuh U, Kürschner H, Battarbee R, et al. Desert plant pollen production and a 160-year record of vegetation and climate change on the Alashan Plateau, NW China[J]. Vegetation History and Archaeobotany, 2006,15(3):181-190. |
| [55] | Ter Braak C, Smilauer P N. CANOCO reference manual and CanoDraw for Windows user’s guide: Software for canonical community ordination (Version 4.5)[M]. Microcomputer Power: Ithaca, 2002. |
| [56] | Bhattacharya T, Kittur S, Sathyakumar S, et al. Diet overlap between wild ungulates and domestic livestock in the greater Himalaya: Implications for management of grazing practices[J]. Proceedings of the Zoological Society, 2012,65(1):11-21. |
| [57] | 苏军虎, Ji Weihong, 徐长林, 等. 甘南草原主要草食动物的食性及其生态位特征[J]. 动物学杂志, 2017,52(3):381-389. |
| [57] | [ Su Junhu, Ji Weihong, Xu Changlin, et al. The ecological characteristics of food habits for four herbivores in Gannan meadow[J]. Chinese Journal of Zoology, 2017,52(3):381-389. ] |
| [58] | 李叶, 王兵, 黄永丽, 等. 天山地区三种自由放牧家畜食性和营养生态位的显微组织分析[J]. 家畜生态学报, 2013,34(9):65-68. |
| [58] | [ Li Ye, Wang Bing, Huang Yongli, et al. Microscopy histological analysis of diet and trophic niche of three free grazing livestock in Tianshan Mountain area[J]. Journal of Domestic Animal Ecology, 2013,34(9):65-68. ] |
| [59] | 江小蕾, 张卫国, 杨振宇, 等. 不同干扰类型对高寒草甸群落结构和植物多样性的影响[J]. 西北植物学报, 2003,23(9):1479-1485. |
| [59] | [ Jiang Xiaolei, Zhang Weiguo, Yang Zhenyu, et al. The influence of disturbance on community structure and plant diversity of alpine meadow[J]. Acta Botanica Boreali-Occidentalia Sinica, 2003,23(9):1479-1485. ] |
| [60] | 仁青吉, 崔现亮, 赵彬彬. 放牧对高寒草甸植物群落结构及生产力的影响[J]. 草业学报, 2008,17(6):134-140. |
| [60] | [ Ren Qingji, Cui Xianliang, Zhang Binbin. Effects of grazing impact on community structure and productivity in an alpine meadow[J]. Acta Prataculturae Sinica, 2008,17(6):134-140. ] |
| [61] | 董全民, 赵新全, 李世雄, 等. 草地放牧系统中土壤-植被系统各因子对放牧响应的研究进展[J]. 生态学杂志, 2014,33(8):2255-2265. |
| [61] | [ Dong Quanmin, Zhao Xinquan, Li Shixiong, et al. A review on the responses of various factors within soil-vegetation system to grazing in grassland grazing systems[J]. Chinese Journal of Ecology, 2014,33(8):2255-2265. ] |
| [62] | 李怡雯, 许清海, 张生瑞, 等. 青藏高原高寒灌丛草甸花粉组合的植被指示性[J]. 科学通报, 2019,64(20):2141-2150. |
| [62] | [ Li Yiwen, Xu Qinghai, Zhang Shengrui, et al. Significance of pollen assemblages for the vegetation composition of alpine shrub meadow in the Qinghai-Tibetan Plateau, China[J]. Chinese Science Bulletin, 2019,64(20):2141-2150. ] |
| [63] | Akeret ?, Haas J N, Leuzinger U, et al. Plant macrofossils and pollen in goat/sheep faeces from the Neolithic lake-shore settlement Arbon Bleiche 3, Switzerland[J]. The Holocene, 1999,9(2):175-182. |
| [64] | Wei H C, Yuan Q, Xu Q H, et al. Assessing the impact of human activities on surface pollen assemblages in Qinghai Lake Basin, China[J]. Journal of Quaternary Science, 2018,33(6):702-712. |
| [65] | 谢端据. 甘青地区史前考古[M]. 北京: 文物出版社, 2002. |
| [65] | [ Xie Duanju. Prehistoric archaeology in Gansu-Qinghai area[M]. Beijing: Cultural Relics Press, 2002. ] |
| [66] | 侯光良, 赖忠平, 孙永娟, 等. 全新世大暖期对青藏高原东北缘人类活动的影响[J]. 干旱区地理, 2013,36(6):971-978. |
| [66] | [ Hou Guangliang, Lai Zhongping, Sun Yongjuan, et al. Impact of the Holocene climatic optimum on human activities in the northeastern margin of the Qinghai-Tibetan Plateau[J]. Arid Land Geography, 2013,36(6):971-978. ] |
| [67] | Chen F H, Dong G H, Zhang D J, et al. Agriculture facilitated permanent human occupation of the Tibetan Plateau after 3600 BP[J]. Science, 2015,347(1):248-250. |
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