横断山南段上新世的植被多样性与分布格局

doi:10.17520/biods.2022295

生物多样性 ›› 2022, Vol. 30 ›› Issue (11): 22295. DOI: 10.17520/biods.2022295

• 中国南方新生代植物多样性演化专题 • 上一篇下一篇

横断山南段上新世的植被多样性与分布格局

黄永江¹^,^*(), 苏涛², 朱海³^,⁴, 贾林波¹, 胡瑾瑾¹, 纪运恒¹, 周浙昆¹^,²

1.中国科学院昆明植物研究所东亚植物多样性与生物地理学重点实验室, 昆明 650201
2.中国科学院西双版纳热带植物园热带森林生态学重点实验室, 云南勐腊 666303
3.上饶师范学院生命科学学院, 江西上饶 334001
4.中国科学院南京地质古生物研究所现代古生物学和地层学国家重点实验室, 南京 210008

收稿日期:2022-06-01 接受日期:2022-08-12 出版日期:2022-11-20 发布日期:2022-08-19
通讯作者: 黄永江
作者简介:* E-mail: huangyongjiang@mail.kib.ac.cn
基金资助:
国家自然科学基金(31700184);国家自然科学基金(41972023);中国科学院南京地质古生物研究所现代古生物学和地层学国家重点实验室开放项目(183132);云南省“万人计划”青年拔尖人才项目(YNWR-QNBJ-2019-261);中国科学院“西部青年学者”人才项目

Vegetation diversity and distribution in the Pliocene of the southern Hengduan Mountains region

Yongjiang Huang¹^,^*(), Tao Su², Hai Zhu³^,⁴, Linbo Jia¹, Jinjin Hu¹, Yunheng Ji¹, Zhekun Zhou¹^,²

1. Key Laboratory for Plant Biodiversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201
2. Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303
3. College of Life Science, Shangrao Normal University, Shangrao, Jiangxi 334001
4. State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008

Received:2022-06-01 Accepted:2022-08-12 Online:2022-11-20 Published:2022-08-19
Contact: Yongjiang Huang

摘要/Abstract

摘要：

横断山南段复杂的地形地貌和多样的气候环境造就了高度丰富的植被多样性。这种植被多样性的演化与形成是植物学、生态学等领域共同关注的科学问题, 而植物化石是回答这一科学问题的重要媒介。本文基于横断山南段的9个上新世化石植物群, 根据其植物组成和优势成分, 分析其所代表的植被类型, 总结了该地区上新世的植被多样性与空间分布, 并结合古环境研究资料, 探讨该地区植被多样性的兴起, 植被分布格局的形成, 及其与环境变迁之间的关联。结果显示, 横断山南段在上新世时期已经具有了多种植被类型, 包括亚热带常绿阔叶林、亚热带落叶阔叶林、硬叶常绿阔叶林、针阔叶混交林、灌丛草地等, 体现了丰富的植被多样性; 植被类型从南面的偏热性植被向北面的偏温性植被逐渐转变, 反映了当时随纬度变化的海拔梯度, 植被类型与分布呈现出与现在高度相似的格局。同时, 小范围内也具有适应于不同气候的植被类型, 反映了当地较大的海拔落差, 可能与高大山体的存在有着密切关联。我们推测, 横断山南段现在丰富的植被多样性和随纬度变化的植被面貌最晚在上新世就已基本形成, 但形成时间是否更早则需要更多、更老化石植物群的发现与研究。

关键词: 横断山, 上新世, 化石植物群, 古植被, 古海拔

Abstract

Aims: Due to complex geomorphology and diverse landscapes, the southern Hengduan Mountains region is home to highly diverse vegetation. The evolution and formation of such diversity is an interesting topic in the fields of botany and ecology, and plant fossils are an important proxy for researching this question.

Methods: In this study, we integrated published sources of nine fossil floras from the Pliocene of the southern Hengduan Mountains region. We first inferred the vegetation types of these fossil floras on the basis of their taxonomic compositions and dominant elements, and then depicted the diversity and spatial distribution of vegetation in the region during the Pliocene. We also correlated the rise of vegetation diversity and the formation of vegetation distribution to major environmental changes in the southern Hengduan Mountains region.

Results: Our synthesis indicates that the southern Hengduan Mountain region already held a variety of vegetation types during the Pliocene, including subtropical evergreen broadleaved forest, subtropical deciduous broadleaved forest, sclerophyllous evergreen broadleaved forest, mixed coniferous broadleaved forest and shrubby meadow. The vegetation tended to have a warm affinity in the south and a cool-temperate affinity in the north, demonstrating altitudinal gradients with latitude at that time. Moreover, vegetation types are shown to be stratified at some localities, suggesting altitudinal gradients at local scales that are likely associated with the occurrence of high mountains. These are highly similar to the modern patterns of vegetation diversity and distribution in this region.

Conclusion: Our study concludes that the modern diversity and distribution of vegetation in the southern Hengduan Mountain region was established no later than the Pliocene. However, more plant fossils from deeper times are needed to verify if the current vegetation was formed at an earlier geological age.

Key words: Hengduan Mountains, Pliocene, fossil flora, palaeovegetation, palaeoelevation

黄永江, 苏涛, 朱海, 贾林波, 胡瑾瑾, 纪运恒, 周浙昆 (2022) 横断山南段上新世的植被多样性与分布格局. 生物多样性, 30, 22295. DOI: 10.17520/biods.2022295.

Yongjiang Huang, Tao Su, Hai Zhu, Linbo Jia, Jinjin Hu, Yunheng Ji, Zhekun Zhou (2022) Vegetation diversity and distribution in the Pliocene of the southern Hengduan Mountains region. Biodiversity Science, 30, 22295. DOI: 10.17520/biods.2022295.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2022295

https://www.biodiversity-science.net/CN/Y2022/V30/I11/22295

图/表 7

图1 横断山南段上新世化石植物群的地理分布。a: 横断山的地理位置与范围; b: 横断山南段上新世化石植物群的空间分布。地图基于GeoMapApp 3.6.10 (Ryan et al, 2009)生成。

Fig. 1 Maps showing the distribution of Pliocene fossil floras in the southern Hengduan Mountains region. a, Geographic position and range of the Hengduan Mountains region; b, Spatial distribution of Pliocene fossil floras in the southern Hengduan Mountains region. The maps were created using GeoMapApp 3.6.10 (Ryan et al, 2009).

表1 横断山南段上新世化石植物群及其地理位置、地层划分与化石类型

Table 1 Pliocene fossil floras in the southern Hengduan Mountains region and their geographic location, stratigraphic horizon and fossil types

植物群名称 Flora	地理位置 Location	经纬度 Geographical coordinate	海拔 Altitude	地层 Stratigraphic horizon	化石类型 Fossil types
庆福植物群 Qingfu Flora	维西县永春乡 Yongchun, Weixi County	99°21′ E, 27°05′ N	2,476 m	三营组 Sanying Formation	叶片 Leaf
红园植物群 Hongyuan Flora	鹤庆县鹤庆盆地西南 Southwestern Heqing Basin, Heqing County	100°08′ E, 26°34′ N	2,291 m	三营组 Sanying Formation	叶片 Leaf
南班榜植物 Nanbanbang Flora	鹤庆县鹤庆盆地西南 Southwestern Heqing Basin, Heqing County	100°10′ E, 26°31′ N	2,200 m	三营组 Sanying Formation	果实、种子 Fruit and seed
福东植物群 Fudong Flora	兰坪县金顶镇 Jinding, Lanping County	99°26′ E, 26°28′ N	2,740 m	三营组 Sanying Formation	叶片、果实、种子、炭屑 Leaf, fruit, seed, and charcoal
三营植物群 Sanying Flora	洱源县三营镇 Sanying, Eryuan County	99°49′ E, 26°00′ N	2,100 m	三营组 Sanying Formation	叶片、孢粉 Leaf and spore/pollen
羊街植物群 Yangjie Flora	永平县永平盆地北 Northern Yongping Basin, Yongping County	99°31′ E, 25°31′ N	1,715 m	三营组 Sanying Formation	叶片 Leaf
羊邑植物群 Yangyi Flora	保山市保山盆地东南 Southeastern Baoshan Basin, Baoshan City	99°15′ E, 24°57′ N	1,650 m	羊邑组 Yangyi Formation	叶片、孢粉 Leaf and spore/pollen
龙陵植物群 Longling Flora	龙陵县龙陵煤矿 Longling Coal Mine, Longling County	98°50′ E, 24°41′ N	1,840 m	羊邑组 Yangyi Formation	孢粉 Spore/pollen
团田植物群 Tuantian Flora	腾冲县团田盆地 Tuantian Basin, Tengchong County	98°38′ E, 24°41′ N	1,250 m	芒棒组 Mangbang Formation	叶片 Leaf

表1 横断山南段上新世化石植物群及其地理位置、地层划分与化石类型

Table 1 Pliocene fossil floras in the southern Hengduan Mountains region and their geographic location, stratigraphic horizon and fossil types

植物群名称 Flora	地理位置 Location	经纬度 Geographical coordinate	海拔 Altitude	地层 Stratigraphic horizon	化石类型 Fossil types
庆福植物群 Qingfu Flora	维西县永春乡 Yongchun, Weixi County	99°21′ E, 27°05′ N	2,476 m	三营组 Sanying Formation	叶片 Leaf
红园植物群 Hongyuan Flora	鹤庆县鹤庆盆地西南 Southwestern Heqing Basin, Heqing County	100°08′ E, 26°34′ N	2,291 m	三营组 Sanying Formation	叶片 Leaf
南班榜植物 Nanbanbang Flora	鹤庆县鹤庆盆地西南 Southwestern Heqing Basin, Heqing County	100°10′ E, 26°31′ N	2,200 m	三营组 Sanying Formation	果实、种子 Fruit and seed
福东植物群 Fudong Flora	兰坪县金顶镇 Jinding, Lanping County	99°26′ E, 26°28′ N	2,740 m	三营组 Sanying Formation	叶片、果实、种子、炭屑 Leaf, fruit, seed, and charcoal
三营植物群 Sanying Flora	洱源县三营镇 Sanying, Eryuan County	99°49′ E, 26°00′ N	2,100 m	三营组 Sanying Formation	叶片、孢粉 Leaf and spore/pollen
羊街植物群 Yangjie Flora	永平县永平盆地北 Northern Yongping Basin, Yongping County	99°31′ E, 25°31′ N	1,715 m	三营组 Sanying Formation	叶片 Leaf
羊邑植物群 Yangyi Flora	保山市保山盆地东南 Southeastern Baoshan Basin, Baoshan City	99°15′ E, 24°57′ N	1,650 m	羊邑组 Yangyi Formation	叶片、孢粉 Leaf and spore/pollen
龙陵植物群 Longling Flora	龙陵县龙陵煤矿 Longling Coal Mine, Longling County	98°50′ E, 24°41′ N	1,840 m	羊邑组 Yangyi Formation	孢粉 Spore/pollen
团田植物群 Tuantian Flora	腾冲县团田盆地 Tuantian Basin, Tengchong County	98°38′ E, 24°41′ N	1,250 m	芒棒组 Mangbang Formation	叶片 Leaf

图2 高山栎组植物的现生和化石叶片(比例尺 = 1 cm)。a: 高山栎组植物的枝条(摄于西藏林芝); b, c: 高山栎组植物现生叶片的正反面及其近叶缘处分叉的二级脉(箭头所示); d-f: 上新世福东植物群的高山栎组叶片化石及其近叶缘处分叉的二级脉(箭头所示)。

Fig. 2 Extant and fossil leaves of Quercus sect. Heterobalanus (scale bars = 1 cm). a, Leafy twigs (picture taken in Linzhi, Xizang); b, c, An extant leaf, showing the upper and lower surfaces, respectively, with the secondary veins bifurcating near the leaf margin (arrows); d-f, Fossil leaves from the Pliocene Fudong flora, with the secondary veins bifurcating near the leaf margin (arrows).

表2 横断山南段上新世化石植物群的优势成分及其所代表的植被类型

Table 2 Dominant elements and vegetation types of the Pliocene fossil floras in the southern Hengduan Mountains region

植物群名称 Flora	优势成分 Dominant element	植被类型 Vegetation type	主要相关文献 Reference
庆福植物群 Qingfu Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2015
红园植物群 Hongyuan Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2016;
南班榜植物群 Nanbanbang Flora	松科、蓼属、悬钩子属、黄芪属、苔草属等 Pinaceae, Polygonum, Rubus, Astragalus, Carex etc.	灌丛草地、针阔叶混交林 Shrubby meadow, mixed coniferous-broadleaved forest	2022
福东植物群 Fudong Flora	高山栎组(叶片); 松科(炭屑) Quercus sect. Heterobalanus (leaves); Pinaceae (charcoals)	硬叶常绿阔叶林(叶片); 针阔叶混交林(炭屑) Sclerophyllous evergreen broadleaved forest (leaves); mixed coniferous-broadleaved forest (charcoals)	2021
三营植物群 Sanying Flora	高山栎组(叶片); 松科、栎属、桦木科等(孢粉) Quercus sect. Heterobalanus (leaves); Pinaceae, Quercus, Betulaceae (spore/pollen)	硬叶常绿阔叶林(叶片); 针阔叶混交林(孢粉) Sclerophyllous evergreen broadleaved forest (leaves); mixed coniferous-broadleaved forest (spore/pollen)	2019
羊街植物群 Yangjie Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2015
羊邑植物群 Yangyi Flora	高山栎组(叶片); 桦木科、壳斗科、金缕梅科等(孢粉) Quercus sect. Heterobalanus (leaves); Betulaceae, Fagaceae, Hamamelidaceae etc. (spore/pollen)	硬叶常绿阔叶林(叶片); 亚热带常绿阔叶林、亚热带落叶阔叶林(孢粉) Sclerophyllous evergreen broadleaved forest (leaves); subtropical evergreen broadleaved forest, subtropical deciduous broadleaved forest (spore/pollen)	2004a
龙陵植物群 Longling Flora	壳斗科、杜鹃花科、冬青科等 Fagaceae, Ericaceae, Aquifoliaceae etc.	亚热带常绿阔叶林 Subtropical evergreen broadleaved forest	2004b
团田植物群 Tuantian Flora	樟科、壳斗科、金缕梅科等 Lauraceae, Fagaceae, Hamamelidaceae etc.	亚热带常绿阔叶林 Subtropical evergreen broadleaved forest	2009

表2 横断山南段上新世化石植物群的优势成分及其所代表的植被类型

Table 2 Dominant elements and vegetation types of the Pliocene fossil floras in the southern Hengduan Mountains region

植物群名称 Flora	优势成分 Dominant element	植被类型 Vegetation type	主要相关文献 Reference
庆福植物群 Qingfu Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2015
红园植物群 Hongyuan Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2016;
南班榜植物群 Nanbanbang Flora	松科、蓼属、悬钩子属、黄芪属、苔草属等 Pinaceae, Polygonum, Rubus, Astragalus, Carex etc.	灌丛草地、针阔叶混交林 Shrubby meadow, mixed coniferous-broadleaved forest	2022
福东植物群 Fudong Flora	高山栎组(叶片); 松科(炭屑) Quercus sect. Heterobalanus (leaves); Pinaceae (charcoals)	硬叶常绿阔叶林(叶片); 针阔叶混交林(炭屑) Sclerophyllous evergreen broadleaved forest (leaves); mixed coniferous-broadleaved forest (charcoals)	2021
三营植物群 Sanying Flora	高山栎组(叶片); 松科、栎属、桦木科等(孢粉) Quercus sect. Heterobalanus (leaves); Pinaceae, Quercus, Betulaceae (spore/pollen)	硬叶常绿阔叶林(叶片); 针阔叶混交林(孢粉) Sclerophyllous evergreen broadleaved forest (leaves); mixed coniferous-broadleaved forest (spore/pollen)	2019
羊街植物群 Yangjie Flora	高山栎组 Quercus sect. Heterobalanus	硬叶常绿阔叶林 Sclerophyllous evergreen broadleaved forest	2015
羊邑植物群 Yangyi Flora	高山栎组(叶片); 桦木科、壳斗科、金缕梅科等(孢粉) Quercus sect. Heterobalanus (leaves); Betulaceae, Fagaceae, Hamamelidaceae etc. (spore/pollen)	硬叶常绿阔叶林(叶片); 亚热带常绿阔叶林、亚热带落叶阔叶林(孢粉) Sclerophyllous evergreen broadleaved forest (leaves); subtropical evergreen broadleaved forest, subtropical deciduous broadleaved forest (spore/pollen)	2004a
龙陵植物群 Longling Flora	壳斗科、杜鹃花科、冬青科等 Fagaceae, Ericaceae, Aquifoliaceae etc.	亚热带常绿阔叶林 Subtropical evergreen broadleaved forest	2004b
团田植物群 Tuantian Flora	樟科、壳斗科、金缕梅科等 Lauraceae, Fagaceae, Hamamelidaceae etc.	亚热带常绿阔叶林 Subtropical evergreen broadleaved forest	2009

图3 上新世南班榜植物群的草本植物果实、种子化石(比例尺 = 0.5 mm)。a: 毛茛属; b: 紫菫属; c: 酸模属; d, e: 蓼属; f: 藜属; g: 繁缕属; h: 草莓属; i: 黄芪属; j: 紫菀属; k: 薹草属; l: 水葱属。b-d, h, j和l来自Huang等(2022), 并获得了出版商的使用许可。

Fig. 3 Fossil fruits and seeds of herbs from the Pliocene Nanbanbang flora (scale bars = 0.5 mm). a, Ranunculus; b, Corydalis; c, Rumex; d, e, Polygonum; f, Chenopodium; g, Stellaria; h, Fragaria; i, Astragalus; j, Aster; k, Carex; l, Schoenoplectus. Images b-d, h, j and l are based on Huang et al (2022) with the formal permission of the publisher.

图4 上新世福东植物群的炭屑化石(比例尺: a-c, e-g, 50 μm; d, h, 10 μm)。a-d: 松柏类; a: 横切面, 显示形态、排列规则的腔隙(木纤维细胞横面观)及木射线(纵面观) (箭头所示); b: 横切面部分放大, 显示形态、排列规则的腔隙及相互联合的细胞壁; c: 纵切面, 显示木纤维细胞(纵面观)、管胞(纵面观)和单列细胞的木射线(横面观); d: 纵切面部分放大, 显示管胞壁上的单列圆形纹孔(箭头所示); e-h: 非松柏类; e: 横切面, 显示不规则的腔隙(导管细胞、木纤维细胞和薄壁细胞); f: 横切面部分放大, 显示导管的横面观(箭头所示); g: 纵切面, 显示单个导管的纵面观; h: 纵切面部分放大, 显示导管细胞壁上不规则排列的纹孔。

Fig. 4 Fossil charcoals from the Pliocene Fudong flora (scale bars = 50 μm (a-c, e-g); 10 μm (d, h)). a-d, Conifers; a, Transverse section, showing the homogenous, regular lumina (xylem cells; transverse view) and xylem rays (longitudinal view) (arrows); b, Amplified view of the transverse section, showing the regular xylem cells and homogenized adjacent cell walls; c, Longitudinal section, showing the xylem cells (longitudinal view), tracheid cells (longitudinal view) and uniseriate xylem rays (transverse view); d, Amplified view of the longitudinal section, showing the uniseriate, circular pits (arrows) on the tracheid wall; e-h, Non-conifers; e, Transverse section, showing the heterogenous, irregular lumina (vessel, xylem and parenchyma); f, Amplified view of the transverse section, showing the transverse view of a vessel (arrow); g, Longitudinal section, showing the longitudinal view of a vessel; h, Amplified view of the longitudinal section, showing alternative pits on the vessel cell wall.

图5 上新世横断山南段的植被分布。地图基于GeoMapApp 3.6.10 (Ryan et al, 2009)生成。

Fig. 5 Vegetation distribution in the Pliocene of the southern Hengduan Mountains region. The map was created using GeoMapApp 3.6.10 (Ryan et al, 2009).

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横断山南段上新世的植被多样性与分布格局

Vegetation diversity and distribution in the Pliocene of the southern Hengduan Mountains region

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