The discovery of Podocarpium from the Miocene of Qaidam Basin and its biogeographic implication
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摘要:
单籽豆属(Podocarpium)是已经灭绝的一类豆科植物, 也是新生代地层中广泛出现的类群。该属在欧亚大陆中有着丰富的化石记录, 但由于缺乏重点区域以及关键时间点的化石证据, 限制了对其演化与分布历史的认识。本研究以青藏高原东北部柴达木盆地瑙格剖面2100 m处的中新世地层中的单籽豆属为研究对象, 结合新生代以来的全球化石记录, 综合概述了该属的生物地理历史, 并依据青藏高原地区的具有准确年代控制的化石记录探讨了环境变化对单籽豆属演变的影响。研究结果表明: 1)依据柴达木盆地中中新世晚期新标本枝叶和荚果的形态比较, 将其定为柄豆荚(Podocarpium podocarpum); 2)依据已知的青藏高原化石记录表明, 单籽豆属在中新世时期的分异度最高且分布特征与全球具有同步性, 同时支持了该属通过低纬度路径扩散和传播的可能性; 3)全球范围内的干旱和降温可能不是导致单籽豆属分布区范围缩小的主要原因, 中新世之后青藏高原部分地区的地形进一步抬升通过改变降水模式促进了该属在亚洲内陆地区分范围的收缩。
Abstract:Podocarpium is an extinct genus of the family Fabaceae that was widely distributed in the Eurasian continent during Cenozoic. But the lack of fossil evidence from key regions and critical time points limits the understanding of its evolutionary and distributional history. In this study, we focus on the Middle Miocene Podocarpium of the Naoge section in the Qaidam Basin, located in the northeastern of the Tibetan Plateau. By integrating global fossil records from the Cenozoic, this paper provides a comprehensive overview of the biogeographical history of the genus and discusses how environmental changes influenced its evolution, based on accurately dated specimens from the Tibetan Plateau. The Naoge section is located about 43 km south of Gahai Town, Delingha City, in the northeastern of the Qaidam Basin, with a thickness of approximately 3300 meters. Exposed are the Upper Cretaceous Quanyagou Formation, the Miocene to Pliocene Lower Youshashan Formation, Upper Youshashan Formation, and the Shizigou Formation. The fossil specimens in this study were collected from the Upper Youshashan Formation of the section, at an elevation of 2100 meters, within greenish-gray muddy siltstone. According to paleomagnetic dating, the geological age of the fossil layer is approximately 11 Ma. Our results show that the well-preserved fossil fruits and leaves of the Podocarpium podocarpum were described from the Middle Miocene of the Qaidam Basin, northeast Tibetan Plateau. Furthermore, fossil records from the Tibetan Plateau indicate that the genus had its highest diversity during the Miocene, with a distribution pattern synchronous with global trends, supporting the possibility that the Podocarpium spread and dispersed via low-latitude routes. Thus, global aridification and cooling are unlikely to be the primary causes of the range contraction of the Podocarpium. Instead, further uplift of certain areas of the Tibetan Plateau after the Miocene, by altering precipitation patterns, which may have contributed to the reduction of the Podocarpium's distribution in inland Asia.
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Key words:
- Miocene /
- Podocarpium /
- biogeography /
- Qaidam Basin
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图 1
单籽豆属(Podocarpium)叶片及荚果形态复原图
Figure 1.
Morphological reconstruction of the Podocarpium, and modified after WGCPC[9])
图 2
单籽豆属(Podocarpium)化石记录的全球地理分布图
Figure 2.
The distribution of megafossil records of Podocarpium throughout the world, and drawing after Table 1
图 3
柴达木盆地瑙格剖面位置及地质概况
Figure 3.
The location of the Naoge section and geological features of the Qaidam Basin. (a)Distribution of study site and fossil records of Podocarpium on the Tibetan Plateau; (b)Paleogeographic location of study site during the Middle Miocene; (c)Geological map of the Naoge(A-B marks the section location); (d)Stratigraphic column of the Naoge section[53]
图 4
柴达木盆地瑙格剖面上油砂山组中的柄豆荚(Podocarpium podocarpum)叶片化石标本
Figure 4.
Leaves of the Podocarpium podocarpum from the Shangyoushashan Formation of Naoge section, Qaidam Basin
图 5
柴达木盆地瑙格剖面上油砂山组中的柄豆荚(Podocarpium podocarpum)荚果化石标本
Figure 5.
Fruits of of the Podocarpium podocarpum from the Shangyoushashan Formation of Naoge section, Qaidam Basin
图 6
地质时期单籽豆属(Podocarpium)的化石点分布(古地理图依据Gplates软件)
Figure 6.
The distribution of fossil sites of the Podocarpium during geological periods, and modified from Gplates
表 1
单籽豆属(Podocarpium) 化石记录(修改自Li等[1]和Han等[2])
Table 1.
Megafossil records of Podocarpium (modified from Li et al. [1] and Han et al. [2])
序号 种名 化石类别 年代 地点 文献 1 Podocarpium eocenicum 荚果 始新世 中国，海南省长昌盆地 [28] 2 Podocarpium eocenicum 荚果 始新世 中国，广东省茂名盆地 [28] 3 Podocarpium tibeticum 荚果&枝叶 始新世 中国，西藏自治区申扎县 [1] 4 Podocarpium podocarpum 荚果&枝叶 早渐新世 中国，柴达木盆地大柴旦 [2] 5 Podocarpium podocarpum 荚果&单叶 渐新世 中国，柴达木盆地花土沟 [29] 6 Podocarpium podocarpum 荚果 渐新世 中国，广西自治区宁明县 [30] 7 Podocarpium podocarpum 枝叶 早渐新世 法国 [29] 8 Podocarpium podocarpum 荚果&枝叶 早渐新世 法国 [20] 9 Podocarpium podocarpum 枝叶 早渐新世 法国 [4] 10 Podocarpium podocarpum 枝叶 晚渐新世 捷克 [4] 11 Podocarpium podocarpum 荚果 中新世 中国，青海省泽库县 [23] 12 Podocarpium podocarpum 荚果 中新世 中国，青海省泽库县 [30] 13 Podocarpium podocarpum 荚果&枝叶 中中新世早期 中国，山东省临朐县山旺镇 [11] 14 Podocarpium podocarpum 荚果&枝叶 早中新世 捷克 [31] 15 Podocarpium podocarpum 荚果&枝叶 早中新世 捷克 [31] 16 Podocarpium podocarpum 单叶 早中新世 捷克 [32] 17 Podocarpium podocarpum 单叶 早中新世 捷克 [4] 18 Podocarpium podocarpum 荚果&枝叶 早中新世 德国 [4] 19 Podocarpium podocarpum / 早中新世 塞尔维亚 [1~2] 20 Podocarpium podocarpum 荚果 早中新世 塞尔维亚 [1~2] 21 Podocarpium podocarpum 荚果&单叶 早中新世 日本，本州岛北部 [33] 22 Podocarpium podocarpum 荚果&枝叶 中中新世 德国 [20] 23 Podocarpium podocarpum 荚果 早-中中新世 匈牙利 [34] 24 Podocarpium podocarpum 单叶 中中新世早期 日本，长崎县 [1~2] 25 Podocarpium podocarpum / 中中新世 意大利 [1~2] 26 Podocarpium podocarpum / 中中新世 德国 [18] 27 Podocarpium podocarpum / 中中新世 德国 [1~2] 28 Podocarpium podocarpum / 中中新世 德国 [1~2] 29 Podocarpium podocarpum / 中中新世 瑞士 [1~2] 30 Podocarpium podocarpum 枝叶 中新世 中国，江苏省泗洪县 [35] 31 Podocarpium podocarpum 枝叶 中新世 中国，江苏省南京市璞真森林农场 [6] 32 Podocarpium podocarpum 枝叶 中新世 中国，江苏省南京市灵岩山 [6] 33 Podocarpium podocarpum 枝叶 中新世 中国，江苏省黄冈市 [6] 34 Podocarpium podocarpum 荚果 中中新世 匈牙利 [1~2] 35 Podocarpium podocarpum 荚果&单叶 中中新世 日本，本州岛中北部 [36] 36 Podocarpium podocarpum 荚果&枝叶 中中新世 中国，青海省德令哈市瑙格地区 当前研究 37 Podocarpium podocarpum 枝叶&单叶 中中新世 日本 [1~2] 38 Podocarpium podocarpum 荚果 中-晚中新世 匈牙利 [1~2] 39 Podocarpium podocarpum 荚果 中中新世晚期 中国，云南开远市 [24] 40 Podocarpium podocarpum 枝叶 中新世 中国，浙江省宁海县 [24] 41 Podocarpium podocarpum 枝叶 中新世 中国，浙江省嵊县 [24] 42 Podocarpium podocarpum / 中-晚中新世 罗马尼亚 [1~2] 43 Podocarpium podocarpum 荚果 中-晚中新世 中国，浙江天台县 [30] 44 Podocarpium podocarpum 枝叶 中-晚中新世 中国，浙江省绍兴市新昌县下屏山 [35] 45 Podocarpium podocarpum 枝叶 中-晚中新世 哈萨克斯坦 [1~2] 46 Podocarpium podocarpum 枝叶 中-晚中新世 波兰 [1~2] 47 Podocarpium podocarpum 荚果&枝叶 晚中新世 德国 [1~2] 48 Podocarpium podocarpum 荚果 晚中新世 瑞士 [4] 49 Podocarpium podocarpum 荚果 晚中新世 瑞士 [1~2] 50 Podocarpium podocarpum / 晚中新世 罗马尼亚 [1~2] 51 Podocarpium podocarpum / 晚中新世 塞尔维亚 [1~2] 52 Podocarpium podocarpum / 晚中新世 意大利 [1~2] 53 Podocarpium podocarpum 荚果&枝叶 晚中新世 德国 [1~2] 54 Podocarpium podocarpum / 晚中新世 罗马尼亚 [1~2] 55 Podocarpium podocarpum 单叶 晚中新世 希腊，克里特岛 [1~2] 56 Podocarpium podocarpum 单叶 中新世 日本，土岐市高野山 [1~2] 57 Podocarpium podocarpum 荚果&单叶 中新世 奥地利 [21] 58 Podocarpium podocarpum 枝叶&单叶 中新世 中国，青海省乌兰县 [24] 59 Podocarpium podocarpum 枝叶&单叶 上新世 中国，江苏省南京市菊花台 [6] 60 Podocarpium podocarpum 枝叶&单叶 上新世 中国，山西省晋中市太谷区 [1] 61 Podocarpium podocarpum 荚果&枝叶 上新世 中国，山西省榆社县 [1] 62 Podocarpium podocarpum 枝叶&单叶 上新世 中国，云南省元谋盆地 [1] 
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