Evolution of Functional Diversity in Terrestrial Vertebrates from the Late Jurassic to Early Cretaceous in Asia
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摘要: 在侏罗纪-白垩纪转折期，全球生态系统经历显著重组，约20%的海洋生物灭绝，陆地上鳄形目、翼龙及四足动物多样性锐减，体型较大的类群受影响尤甚. 然而，该时期陆地生态多样性的演化仍缺乏系统约束. 亚洲地区保存连续的晚侏罗世至早白垩世地层，脊椎动物化石丰富，为重建该转折期陆地生态演化提供理想材料. 收集亚洲脊椎动物出现记录，结合生活习性、食性与体型信息建立生态分类体系，并通过体型量化与重采样校正采样偏差. 结果显示，大型蜥臀目恐龙（下降50%~80%）、龟类（40%~50%）和哺乳动物（60%~70%）多样性显著下降，而淡水鱼类及部分爬行动物受影响较小. 生态空间分析表明，该转折期亚洲陆地生态系统在物种多样性与功能结构上均发生显著调整. 此外，相关性分析表明植被更替促进了水生类群多样性上升，而气候变暖与干旱化趋势对陆生类群多样性产生明显抑制作用.Abstract: During the Jurassic-Cretaceous (J-K) transition, global ecosystems underwent profound changes. Studies on biodiversity during this period indicate that approximately 20% of marine species went extinct in shallow-sea environments, while on land, crocodyliform diversity declined by 55%-75%, and tetrapods and pterosaurs experienced a 75%-80% reduction in diversity. Overall, larger-bodied taxa were disproportionately affected. However, the evolutionary trajectory of terrestrial ecological diversity during this pivotal transition remains poorly constrained.In Asia, stratigraphic successions spanning the Late Jurassic to Early Cretaceous are well developed and have yielded abundant vertebrate fossils, making the region a key area for reconstructing the evolutionary history of terrestrial ecological diversity across the J-K transition. In this study, we compiled occurrence data of vertebrate fossils from the Late Jurassic to Early Cretaceous of Asia and integrated species-level ecological traits for analysis. Ecological classification was established based on habitat, diet, and body size, with body size data measured to refine trait differentiation. Resampling methods were applied to correct for sampling bias and uneven sample sizes.The results show that large-bodied saurischian dinosaurs (by approximately 50%-80%), turtles (about 40%-50%), and mammals (about 60%-70%) experienced marked declines in diversity across the J-K boundary, whereas freshwater fishes and some other reptilian groups were less affected. Analyses of ecospace structure reveal substantial adjustments in both species diversity and functional structure within Asian terrestrial ecosystems during the J-K transition. Furthermore, responses to environmental factors varied markedly among clades: overall, vegetation changes appear to have promoted increases in aquatic diversity, whereas climatic warming and increasing aridity exerted strong suppressive effects on the diversity of terrestrial groups.
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Key words:
- Early Cretaceous /
- taxonomic diversity /
- functional diversity /
- ecospace /
- biota /
- geobiology
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图 1 晚侏罗世至早白垩世亚洲地区陆地脊椎动物重采样物种多样性变化
a.亚洲地区所有陆地脊椎动物出现点数据重采样物种多样性（q=0.3）；b.翼龙重采样物种多样性；c.蜥臀目（方形）和鸟臀目（圆形）重采样物种多样性；d.鳄形目（菱形）和哺乳动物（三角形）重采样物种多样性；e.两栖类（圆形）和龟类（方形）重采样物种多样性；f.软骨鱼类（圆形）和其它鱼类（十字形）重采样物种多样性；黑色实线代表所有出现点数据重采样结果，紫色虚线代表排除热河、燕辽生物群后重采样结果；Baj. 巴柔阶；Bat. 巴通阶；Ca. 卡洛夫阶；Ox. 牛津阶；Ki. 钦莫利阶；Ti. 提塘阶；Be. 贝里阿斯阶；Va. 瓦兰今阶；Ha. 欧特里夫阶；Bar. 巴雷姆阶；Apt. 阿普特阶；Alb. 阿尔布阶
Fig. 1. Subsampled diversity of terrestrial vertebrates in Asia from the Late Jurassic to the Early Cretaceous
图 2 晚侏罗世至早白垩世亚洲地区陆地脊椎动物原始数据与重采样后物种多样性与新生率、灭绝率变化
a~c.使用原始数据所得物种多样性与新生率、灭绝率变化；d~f.重采样（q=0.3）后所得物种多样性与新生率、灭绝率变化，淡色区域为95%置信区间；a，d. 基于延续范围（橙色实线；假设物种在其首次和末次出现之间连续存在，即使中间阶段没有采样到，也被视为仍存在）与矫正采样分箱（绿色实线；仅统计在特定时间箱中实际采集到的化石种类，但通过统计方法对采样量差异进行校正）的物种多样性曲线；b，e. 使用per-capita方法计算的灭绝率与新生率. 红色为灭绝率，蓝色为新生率；c，f. 使用second-for-third方法计算的灭绝率与新生率；红色为灭绝率，蓝色为新生率；Aal. 阿林阶；Baj. 巴柔阶；Bat. 巴通阶；Ca. 卡洛夫阶；Ox. 牛津阶；Ki. 钦莫利阶；Ti. 提塘阶；Be. 贝里阿斯阶；Va. 瓦兰今阶；Ha. 欧特里夫阶；Bar. 巴雷姆阶；Apt. 阿普特阶；Alb. 阿尔布阶
Fig. 2. Changes in species diversity, speciation and extinction rate of Asian terrestrial vertebrates from the Late Jurassic to the Early Cretaceous based on raw and subsampled data
图 3 晚侏罗世至早白垩世亚洲地区陆地生物空间格子与各功能群相对含量
a.功能群相对含量，以出现点数量为计数；b.功能群相对含量，以属数量为计数；c. 空间格子分布随时间变化，以出现点数量为计数；Aal. 阿林阶；Baj. 巴柔阶；Bat. 巴通阶；Ca. 卡洛夫阶；Ox. 牛津阶；Ki. 钦莫利阶；Ti. 提塘阶；Be. 贝里阿斯阶；Va. 瓦兰今阶；Ha. 欧特里夫阶；Bar. 巴雷姆阶；Apt. 阿普特阶；Alb. 阿尔布阶
Fig. 3. Relative abundanceand ecospace cubesof functional groups of Asian terrestrial species from the Late Jurassic to the Early Cretaceous
图 4 晚侏罗世至早白垩世亚洲地区地层数以及采集数和出现点数分布
Aal. 阿林阶；Baj. 巴柔阶；Bat. 巴通阶；Ca. 卡洛夫阶；Ox. 牛津阶；Ki. 钦莫利阶；Ti. 提塘阶；Be. 贝里阿斯阶；Va. 瓦兰今阶；Ha. 欧特里夫阶；Bar. 巴雷姆阶；Apt. 阿普特阶；Alb. 阿尔布阶
Fig. 4. Distribution of the count of formations, collections, and occurrences of Asian terrestrial vertebrates from the Late Jurassic to the Early Cretaceous
表 1 陆地生物生态特征划分标准
Table 1. Classification Criteria for Ecological Characteristics of Terrestrial Organisms
生态特征 描述 例子 生活习性（tiering） 1-水生（aquatic） 在水中繁殖，生活 鱼类 2-两栖（amphibious） 能短暂离开水体，但依然依靠水体繁殖 肺鱼和两栖类 3-穴居（troglobitic） 在地下生活（例如洞穴） 少数哺乳动物 4-地面生活（ground dwelling） 地面上生活 大部分爬行动物，部分哺乳动物 5-树上生活（arboreal） 大部分时间在高植被上生活 大部分哺乳动物 6-空中生活（aerial） 大量时间在空中，且需要利用飞行完成日常活动 鸟类、翼龙 食性（diet） 1-滤食性（filter feeding） 通过滤食水体中的食物颗粒存活 翼龙 2-植食性（herbivore） 以植物作为主要能量来源 部分哺乳动物，爬行类 3-杂食性（omnivore） 以动物组织，植物为食 部分哺乳动物，爬行类 4-肉食性（carnivore） 通过腐食、捕食到的动物组织作为主要能量来源 爬行类，哺乳动物，鱼类 体型（size of body） 1-非常小（tiny） < 50 cm 鱼类，哺乳动物，青蛙 2-很小（very small） 50~100 cm 部分爬行类，哺乳动物 3-小（small） 101~200 cm 部分爬行类 4-中（medium） 201~500 cm 部分爬行类 5-大（large） 501~800 cm 爬行类，以恐龙为主 6-很大（very large） 801~1 500 cm 爬行类，以恐龙为主 7-非常大（extremely large） > 1 500 cm 爬行类，以蜥臀目恐龙为主 
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表 2 各门类生物多样性与环境因子之间的模型拟合关系
Table 2. Biodiversity-environment relationships modeled for different taxonomic group
种群 环境因子 Spearman Pearson AICc Rho P值 r P值 AICc 权重 对数似然 翼龙 海平面变化 0.561 0.116 0.497 0.173 67.589 0.152 -28.395 鸟臀目 温度 -0.597 0.090 -0.781 0.013 52.747 0.716 -20.973 蜥臀目 裸子植物 -0.817 0.011 -0.772 0.015 67.555 0.493 -28.378 温度 -0.717 0.037 -0.743 0.022 68.469 0.312 -28.834 哺乳类 温度 -0.496 0.175 -0.608 0.082 60.137 0.296 -24.668 鳄形目 裸子植物 0.862 0.003 0.944 0.000 35.387 0.949 -12.293 龟类 二氧化碳浓度 -0.644 0.061 -0.660 0.053 45.781 0.271 -17.490 亚洲大陆极移 -0.610 0.108 -0.572 0.138 44.404 0.539 -16.202 鱼类 裸子植物 0.812 0.008 0.895 0.001 54.255 0.809 -21.728 软骨鱼 被子植物 0.753 0.019 0.659 0.053 36.874 0.301 -13.037 地层数 0.596 0.090 0.645 0.061 37.170 0.260 -13.185 裸子植物 0.596 0.090 0.621 0.074 37.624 0.207 -13.412
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