The discovery of the Eocene genus Palibinia from Xizang, China and its geological and biological significances
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摘要:
本研究重新研究了发现于西藏班戈盆地牛堡组、最初被定为似班克木属(cf. Banksia)的叶化石标本。研究结果表明: 这些标本的叶形、叶片大小和叶脉特征均与狭叶梅属(Palibinia)的欧亚狭叶梅(Palibinia laxifolia Korovin)一致, 因此将这些标本修订为欧亚狭叶梅。狭叶梅属是一类已经灭绝的植物, 依据目前所保存及可观察到的形态特征, 还不能确定狭叶梅属与现代植物类群的亲缘关系。化石记录显示: 狭叶梅属最早出现于广东三水、湖南衡阳、江苏仪征等地的古新世地层, 在始新世狭叶梅属的分布区进一步扩大, 中始新世狭叶梅属出现在青藏高原、中亚、欧洲和北美, 渐新世狭叶梅属的分布区进一步缩小, 在晚渐新世以后完全消失。此前的研究从狭叶梅属的形态特征推论, 该属是干旱环境的指示植物。本研究对全球狭叶梅属分布的24个化石产地古气候模拟数据进行了分析, 结果表明, 狭叶梅属是一类生态幅较为广泛的植物, 基于西藏蒋浪和四川热鲁中始新世植物群重建的古气候结果, 也进一步证明狭叶梅属可分布于湿润环境, 不宜作为干旱环境的指示植物。古气候模型模拟还表明狭叶梅属分布的区域年均温都比较高, 是一类适应于热室气候环境的植物, 属于古近纪中国东南陆地生态系统的重要类群。
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关键词:
- 狭叶梅属 /
- 古近纪 /
- 干旱环境 /
- 陆地生态系统重要类群 /
- 热室气候
Abstract:The genus Palibinia Korovin is an extinct group of angiosperms that was widely distributed across China during the Paleogene, with records also found in center Asia, Europe and America from the Middle Eocene to the Late Oligocene. It was initially recognized as an indicator of arid environments based on its morphological traits and geographical distribution, but recent studies have challenged this assumption.
This research analyzed leaf fossil specimens previously identified as "cf. Banksia leaf" from the Jianglang flora in the Niubao Formation, Bangor Basin, Xizang. The study applied the HadCM3 climate model, developed by the University of Bristol, to simulate paleoclimates over five periods from the Paleocene to the Oligocene. The paleoclimate for the 24 Palibinia fossil sites were extracted from these simulations. Additionally, paleoclimates for the Jianglang flora in the Niubao Formation and the Relu flora in the Relu Formation (Middle Eocene) were reconstructed using existing data.
The Jianglang flora (31.63°N, 90.03°E; 4850 m a.s. l.) is located in the Bangor Basin, near the Bangong-Nujiang Suture Zone of e central Tibetan Plateau. The Niubao Formation is preliminarily dated to the Late Paleocene-Early Eocene, with fossils embedded in lacustrine mudstones and siltstones. In an approximately 32 m thick outcrop, five layers of well-preserved fossils, including plants, mammals, fishes, and insects, have been discovered. Detrital zircon U-Pb dating, along with floristic comparison, placed the age of Jianglang flora within the Middle Eocene, around 47 Ma.
The results revealed that the leaf shape, size, and venation pattern of "cf. Banksia leaf" specimens closely resemble Palibinia laxifolia Korovin, leading to their reclassification. Despite thorough the detailed morphological analysis, the phylogenetic relationship of Palibinia to modern groups remains undetermined. According to fossil records, Palibinia initially appeared in the Paleocene strata of Sanshui (Guangdong Province), Hengyang (Hunan Province), and Yizheng (Jiangsu Province). During the Middle Eocene, its distribution expanded to the Qinghai-Tibetan Plateau, Central Asia, Europe, and North America. However, by the Late Eocene, its range contracted, eventually culminating in its extinction during the Late Oligocene. Palibinia played a crucial role in the terrestrial ecosystems of Southeast China during the Paleogene. Although previous studies linked that Palibinia to arid conditions based on its morphological features, our simulations of paleoclimate data from 24 global fossil sites suggest that Palibinia occupied a broad ecological niche. Paleoclimate reconstructions of the Middle Eocene fossil assemblages from the Jianglang flora (Xizang) and the Relu flora (Sichuan) demonstrate that Palibinia also thrived in humid environments, challenging its classification as an arid indicator. Furthermore, paleoclimate stimulations show that regions inhabited by Palibinia typically experienced high average annual temperatures, signifying its adaptation to hothouse climates. As the global climate shifted towards cooler conditions since the Late Eocene, the distribution of Palibinia declined, culminating in its worldwide extinction globally during the Late Oligocene.
Based on the research conducted, the following conclusions can be drawn: (1) Palibinia laxifolia Korovin was a crucial component of terrestrial ecosystems in eastern Asia during the Paleogene and was well-adapted to a hothouse climate; (2) Palibinia laxifolia likely originated in southeastern China and subsequently spread to other regions of the Northern Hemisphere; (3) The discovery of Palibinia laxifolia in the Middle Eocene strata of the Bangor Basin in Tibet indicates that there was floristic exchange between the Qinghai-Tibet Plateau and southeastern China during the Paleogene; (4) Palibinia laxifolia exhibited a broad ecological niche and thrived in various environments, suggesting that it cannot be considered an indicator of arid conditions.
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Key words:
- Palibinia /
- Paleogene /
- arid environment /
- terrestrial ecosystem /
- hothouse climate
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图 2
蒋浪化石点位置(a)及地质概况简图(b)
Figure 2.
Location of the Jianglang fossil site(a)and simplified geological map(b)
图 3
产自西藏班戈牛堡组的欧亚狭叶梅Palibinia laxifolia Korovin叶片照片
Figure 3.
Leaves of Palibinia laxifolia Korovin from the Niubao Formation of Bange basin of Xizang
图 4
欧亚狭叶梅Palibinia laxifolia Korovin叶片形态细节
Figure 4.
Details of leaf morphology of Palibinia laxifolia Korovin. In Fig. 4, 1 is the main pulse, 2 is the secondary pulse, and 3 is the secondary pulse branch; scale bar=1 mm
图 5
不同时代24个狭叶梅属化石点古气候差异的箱型图
Figure 5.
Box plot of paleoclimate differences at 24 fossil sites in different geological ages. Box plots illustrating the paleoclimate are(a)annual precipitation, (b)precipitation of coldest quarter, (c)precipitation of warmest quarter, (d)average annual temperature, (e)mean temperature of coldest quarter, and(f)mean temperature of warmest quarter in different periods. The p-values displayed above each figure represent the results of ANOVA test, which assesses whether there are significant differences in the mean values of paleoclimate factors across the five geological periods. A p-value of ≤ 0.05 indicates a significant difference; a p-value of ≤ 0.01 indicates a highly significant difference; a p-value of ≤ 0.001 indicates an extremely significant difference; and a p-value of ≥ 0.05 indicates no significant difference. In addition, each figure also illustrates the differences in paleoclimate factors between pairs of geological periods. The number of asterisks(*)indicates the significance level of the differences: *: p < =0.05, **: p < =0.01, ***: p < =0.001, ns: p≥0.05. The boxes represent the 25th and 75th percentiles, the horizontal line represents the median, the whiskers represent the 5th to 95th percentiles, and the circles represent outliers
图 6
产自湖南茶山坳古新统(a，b，d，e)和广东南海盐布(c)始新统的欧亚狭叶梅
Figure 6.
Leaves of Palibinia laxifolia Korovin from the Paleocene of Chashanao, Hunan(a, b, d, e)and the Eocene of Yanbu, Guangdong(c); scale bar=1 cm
表 1
狭叶梅属分布点及古气候模拟数据信息
Table 1.
Global distribution of Palibina and their paleoclimate simulation
编号 化石点 国家/地区 时代 纬度
(°)经度
(°)年均温
(℃)夏季均温
(℃)冬季均温
(℃)年降水
(mm)夏季降水
(mm)冬季降水
(mm)参考文献 1 大仪集，阜宁组 中国，江苏 古新世 32.55 119.25 19.6 28.0 11.7 1332 439 301 [2] 2 清江，清江组下段 中国，江西 古新世 28.05 115.29 21.8 30.2 12.6 920 346 131 [2] 3 三水，布心组 中国，广东 古新世 23.22 113.02 22.8 29.6 14.1 1398 663 98 [4] 4 衡阳，茶山坳组 中国，湖南 古新世 26.93 112.68 20.3 29.7 9.2 1153 490 113 [10] 5 当阳，方家河组 中国，湖北 古新世 30.80 111.79 20.4 29.2 12.0 519 198 90 [2] 6 蒋浪，牛堡组 中国，西藏 中始新世 31.63 90.03 22.6 40.7 4.7 235 127 6 [19] 7 理塘，热鲁组 中国，四川 中始新世 29.59 100.19 19.8 34.5 4.5 531 265 21 [1] 8 绿河组 美国，犹他 中始新世 39.90 -109.16 20.7 35.7 8.8 429 65 123 [16] 9 绿河组 美国，科罗拉多 中始新世 39.48 -108.60 20.8 35.8 8.9 479 66 147 [16] 10 临江，临江组 中国，江西 晚始新世 28.01 115.41 27.2 39.6 11.5 866 375 22 [2] 11 巴德黑兹 土库曼斯坦 晚始新世 35.65 62.28 26.2 46.5 7.7 91 34 10 [12] 12 艾尔-奥伊兰杜斯 土库曼斯坦 晚始新世 35.80 61.46 26.4 46.8 7.7 87 31 10 [12] 13 Tortmola植物群 哈萨克斯坦 晚始新世 48.55 65.70 17.5 41.3 -2.2 359 46 143 [31] 14 吴城，五里堆组 中国，河南 晚始新世 32.43 113.46 24.5 41.2 6.6 815 343 18 [5] 15 本布里奇 英国，怀特岛 晚始新世 50.74 -1.35 19.5 35.7 8.7 1078 54 460 [15] 16 张易，大坪沟 中国，宁夏 始新世 35.82 106.10 15.4 33.5 -1.8 646 247 50 [3] 17 衡阳，鱼子塘 中国，湖南 始新世 26.89 111.77 27.0 40.0 11.8 710 328 24 [3] 18 桃源，剪家溪组 中国，湖南 始新世 28.89 111.44 26.3 40.6 10.2 597 316 20 [3] 19 天水，固原群 中国，甘肃 始新世 37.16 106.79 14.7 33.5 -3.2 725 254 75 [3] 20 湘乡，下湾铺组 中国，湖南 始新世 27.93 112.59 26.6 40.1 11.0 676 340 26 [3] 21 濮阳 中国，河南 始新世 35.75 115.00 21.4 40.0 2.6 841 346 49 [8] 22 泰安，大汶口 中国，山东 始新世 35.59 117.07 20.9 39.4 1.5 879 356 61 [2] 23 渭南，白鹿塬组 中国，陕西 渐新世 34.33 109.54 14.5 32.2 -3.2 428 232 4 [9] 24 罗特 德国，慕尼黑 晚渐新世 50.76 -7.27 15.0 24.6 8.4 1267 124 453 [13] 
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