更新世中国季风区林栖哺乳动物的森林动态代用指标

doi:10.19615/j.cnki.2096-9899.250311

摘要/Abstract

摘要：

季风对植被的发育具有重要的影响，进而对以植物为食的动物群的演化产生显著的作用。由于素食动物的多样性与植被协同演化，因此可以将之作为森林或植被动态的代用指标。选择了36个具有代表性的产于中国季风区的更新世哺乳动物群作为研究材料，用不同的方法统计了这些动物群中林栖种类和素食种类的多样性，并计算了这些动物群在各个植物分区的合意梯度系数。结果显示，从早更新世到晚更新世，随着东亚夏季和冬季季风的不断交替和加强，森林的活力逐渐波动式下降；而草原的活力不断波动式增加，尤其是在秦淮线以北的地区。因此这些动物群多样性的分析可以作为史前森林动态的代用指标之一。此外，这些代用指标与黄土-古土壤序列和深海氧同位素阶段的对比可以帮助提高生物年代对比的精度及古环境重建。

关键词: 素食类, 动物群, 多样性, 古环境重建, 植物分区

Abstract:

Monsoon has an important impact on the development of vegetation that subsequently has significant influence on the evolution of plant consumers. The diversities of forest dwellers or herbivores follow the evolution of the vegetation, and it is therefore possible to take such diversities as forest or vegetation dynamic proxies. The present work selected 36 Pleistocene faunas of large mammals from monsoon-dominated provinces in China as materials and calculated the diversities of forest dwellers and herbivores with different approaches, as well as the consensus gradient coefficients of all the selected faunas in different flora regionalized subkingdoms. The results show that with the evolution and transitions of the East Asian summer and winter monsoon intensities, the forest vitality decreased while steppe vitality increased gradually in a fluctuated way from the Early Pleistocene to the Late Pleistocene, especially in the provinces north of the Qinling-Huaihe Line. The analyses of such diversities of the faunas can help to determine the forest dynamic proxies. Moreover, the correlation of such proxies to loess-paleosol sequences and marine isotope stages can in turn help to improve the accuracy of dating fauna ages and paleoenvironment reconstruction.

Key words: herbivore, fauna, diversity, paleoenvironment reconstruction, flora regionalization

中图分类号:

Q915.5

白炜鹏, 董为, 张立民, 刘文晖. 更新世中国季风区林栖哺乳动物的森林动态代用指标. 古脊椎动物学报, 2025, 63(2): 133-158.

BAI Wei-Peng, DONG Wei, ZHANG Li-Min, LIU Wen-Hui. The Pleistocene mammalian forest dwellers in monsoon dominated provinces of China as forest dynamics proxies. Vertebrata Palasiatica, 2025, 63(2): 133-158.

图/表 7

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Code	Fauna	Source	Abbreviation	Age	Source
A	Upper Cave, Zhoukoudian	Pei, 1940	SDD*	18-27 ka	Dong and Zhang, 2022
B	Yanjiagang, Harbin	Northeastern Expedition Team of Heilongjiang Managing Committee for Cultural Relics et al., 1987	YJG	21-33 ka	Wei et al., 1986
C	Gulongshan, Dalian	Zhou et al., 1990	GLS	27-30 ka	Dong and Zhang, 2022
D	Shanchengzi, Benxi	Zhang et al., 1986	SCZ	30 ka	Dong and Zhang, 2022
E	Yushu, Yushu	Institute of Vertebrate Paleontology, 1959	YS	27-40 ka	Sun et al., 1981
F	Xiaogushan, Haicheng	Dong et al., 2010	XGS	30-80 ka	Dong and Zhang, 2022
G	Salawusu, Ordos	Qi, 1975; Tong et al., 2008	SLWS	61-68 ka	Yin and Huang, 2004
H	Fuyan Cave, Daoxian	Liu et al., 2015	FYD	80-120 ka	Liu et al., 2015
I	Lingjing, Xuchang	Li and Dong, 2007; Zhang et al., 2009	LJ	105-125 ka	Li et al., 2017
J	Zhiren Cave, Chongzuo	Jin et al., 2009a	MLS*	130-190 ka	Ge et al., 2020
K	Dingcun, Xiangfen	Pei, 1958	DC	160-210 ka	Li et al., 2024
L	Hualongdong, Dongzhi	Wu et al., 2019	Hld	275-331 ka	Wu et al., 2019
M	Xinghuashan, Nanzhao	Qiu et al., 1982	Xhs	150-400 ka	Dong et al., 2020
A	Layers 3-5 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd U^#	230-450 ka	Dong and Zhang, 2022
D	Miaohoushan, Benxi	Zhang et al., 1986	MHS	200-500 ka	Zhang et al., 2007
N	Qizianshan, Yiyuan	Lü et al., 1989	Qzas	320-420 ka	Han et al., 2016
O	Huloudong, Tangshan	Wu et al., 2002	Ts	350-500 ka	Chen et al., 1996; Wang et al., 2002
A	Layers 6-7 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd M^#	450-460 ka	Dong and Zhang, 2022
A	Layers 8-11 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd L^#	460-770 ka	Dong and Zhang, 2022
P	Chenjiawo, Lantian	Qi, 1989	Cjw	650 ka	An et al., 1990a
Q	Donghe, Luonan	Xue et al., 1999	Dh	500-700 ka	Dong et al., 2020
R	Longtandong, Hexian	Huang et al., 2012	Ltd	490-800 ka	Dong and Zhang, 2022
S	Bailongdong, Yunxi	Wu et al, 2009	Bld	500-850 ka	Dong et al., 2020
T	Meipu, Yunxian	Xu, 1978	Mp	500-850 ka	Dong et al., 2020
U	Quyuanhekou, Yunxian	Ji, 1999	Qyhk	936 ka	de Lumley et al., 2008
V	Dadong, Chongzuo	Jin et al., 2009b	DD	0.6-1.2 Ma	Dong and Zhang, 2022
W	Gongwangling, Lantian	Hu and Qi, 1978	Gwl	1.1-1.3 Ma	Dong and Zhang, 2022
X	Juyuandong, Liucheng	Pei, 1962, 1987; Han, 1987; Han and Xu, 1989	JYD	1.15-1.5 Ma	Dong and Zhang, 2022
Y	Longgudong, Jianshi	Zheng, 2004	Lgd	1.3-1.6 Ma	Dong and Zhang, 2022
Z	Mohui, Tiandong	Wang et al., 2007	Mh	1.2-1.8 Ma	Dong et al., 2020
a	Danawu, Yuanmou	Lin et al., 1978	YmM*	1.4-1.8 Ma	Dong and Zhang, 2022
b	Longgupo, Wushan	Huang et al., 1991	LGP	1.5-1.9 Ma	Dong and Zhang, 2022
d	Chutoulang, Chifeng	Zhang, 1989; You and Zhang, 1989; Dong et al., 2017	CTL	1.6-1.9 Ma	Dong et al., 2020
e	Renzidong, Fanchang	Jin and Liu, 2009	RZD	1.7-2.1 Ma	Dong and Zhang, 2022
f	Nihewan, Yangyuan	Teilhard de Chardin and Piveteau, 1930; Qiu, 2000; Tong et al., 2011b	NHW	1.75-2.2 Ma	Dong and Zhang, 2022
g	Longdan, Linxia	Qiu et al., 2004; Deng, 2012	LD	2.16-2.55 Ma	Qiu et al., 2004

Code	Fauna	Source	Abbreviation	Age	Source
A	Upper Cave, Zhoukoudian	Pei, 1940	SDD*	18-27 ka	Dong and Zhang, 2022
B	Yanjiagang, Harbin	Northeastern Expedition Team of Heilongjiang Managing Committee for Cultural Relics et al., 1987	YJG	21-33 ka	Wei et al., 1986
C	Gulongshan, Dalian	Zhou et al., 1990	GLS	27-30 ka	Dong and Zhang, 2022
D	Shanchengzi, Benxi	Zhang et al., 1986	SCZ	30 ka	Dong and Zhang, 2022
E	Yushu, Yushu	Institute of Vertebrate Paleontology, 1959	YS	27-40 ka	Sun et al., 1981
F	Xiaogushan, Haicheng	Dong et al., 2010	XGS	30-80 ka	Dong and Zhang, 2022
G	Salawusu, Ordos	Qi, 1975; Tong et al., 2008	SLWS	61-68 ka	Yin and Huang, 2004
H	Fuyan Cave, Daoxian	Liu et al., 2015	FYD	80-120 ka	Liu et al., 2015
I	Lingjing, Xuchang	Li and Dong, 2007; Zhang et al., 2009	LJ	105-125 ka	Li et al., 2017
J	Zhiren Cave, Chongzuo	Jin et al., 2009a	MLS*	130-190 ka	Ge et al., 2020
K	Dingcun, Xiangfen	Pei, 1958	DC	160-210 ka	Li et al., 2024
L	Hualongdong, Dongzhi	Wu et al., 2019	Hld	275-331 ka	Wu et al., 2019
M	Xinghuashan, Nanzhao	Qiu et al., 1982	Xhs	150-400 ka	Dong et al., 2020
A	Layers 3-5 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd U^#	230-450 ka	Dong and Zhang, 2022
D	Miaohoushan, Benxi	Zhang et al., 1986	MHS	200-500 ka	Zhang et al., 2007
N	Qizianshan, Yiyuan	Lü et al., 1989	Qzas	320-420 ka	Han et al., 2016
O	Huloudong, Tangshan	Wu et al., 2002	Ts	350-500 ka	Chen et al., 1996; Wang et al., 2002
A	Layers 6-7 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd M^#	450-460 ka	Dong and Zhang, 2022
A	Layers 8-11 of Loc. 1, Zhoukoudian	Li and Ji, 1981	Zkd L^#	460-770 ka	Dong and Zhang, 2022
P	Chenjiawo, Lantian	Qi, 1989	Cjw	650 ka	An et al., 1990a
Q	Donghe, Luonan	Xue et al., 1999	Dh	500-700 ka	Dong et al., 2020
R	Longtandong, Hexian	Huang et al., 2012	Ltd	490-800 ka	Dong and Zhang, 2022
S	Bailongdong, Yunxi	Wu et al, 2009	Bld	500-850 ka	Dong et al., 2020
T	Meipu, Yunxian	Xu, 1978	Mp	500-850 ka	Dong et al., 2020
U	Quyuanhekou, Yunxian	Ji, 1999	Qyhk	936 ka	de Lumley et al., 2008
V	Dadong, Chongzuo	Jin et al., 2009b	DD	0.6-1.2 Ma	Dong and Zhang, 2022
W	Gongwangling, Lantian	Hu and Qi, 1978	Gwl	1.1-1.3 Ma	Dong and Zhang, 2022
X	Juyuandong, Liucheng	Pei, 1962, 1987; Han, 1987; Han and Xu, 1989	JYD	1.15-1.5 Ma	Dong and Zhang, 2022
Y	Longgudong, Jianshi	Zheng, 2004	Lgd	1.3-1.6 Ma	Dong and Zhang, 2022
Z	Mohui, Tiandong	Wang et al., 2007	Mh	1.2-1.8 Ma	Dong et al., 2020
a	Danawu, Yuanmou	Lin et al., 1978	YmM*	1.4-1.8 Ma	Dong and Zhang, 2022
b	Longgupo, Wushan	Huang et al., 1991	LGP	1.5-1.9 Ma	Dong and Zhang, 2022
d	Chutoulang, Chifeng	Zhang, 1989; You and Zhang, 1989; Dong et al., 2017	CTL	1.6-1.9 Ma	Dong et al., 2020
e	Renzidong, Fanchang	Jin and Liu, 2009	RZD	1.7-2.1 Ma	Dong and Zhang, 2022
f	Nihewan, Yangyuan	Teilhard de Chardin and Piveteau, 1930; Qiu, 2000; Tong et al., 2011b	NHW	1.75-2.2 Ma	Dong and Zhang, 2022
g	Longdan, Linxia	Qiu et al., 2004; Deng, 2012	LD	2.16-2.55 Ma	Qiu et al., 2004

Fauna*	Total taxa	Forest dwellers	Mixed dwellers	Grassland dwellers	Forest dweller (%)	BSC-PPFF
5 SDD	25	12	2	11	48.00	89.50
3 YJG	21	6	2	13	28.57	80.50
5 GLS	34	15	3	16	44.12	86.00
5 SCZ	27	14	1	12	51.85	91.50
3 YS	25	10	1	14	40.00	83.50
5 XGS	33	15	3	15	45.45	90.00
2 SLWS	18	5	1	12	27.78	81.00
5 FYD	27	20	3	4	74.07	100.50
5 LJ	16	8	2	6	50.00	90.00
7 MLS	20	16	2	2	80.00	97.50
5 DC	16	7	2	7	43.75	88.00
5 Hld	11	8	2	1	72.73	92.75
5 Xhs	19	11	2	6	57.89	89.50
5 Zkd U	33	19	2	12	57.58	93.50
5 MHS	21	14	0	7	66.67	97.00
5 Qzas	13	8	1	4	61.54	93.25
5 Ts	17	10	3	4	58.82	94.50
5 Zkd M	21	11	3	7	52.38	92.00
5 Zkd L	36	18	4	14	50.00	89.75
5 Cjw	8	6	1	1	75.00	92.00
5 Dh	18	11	2	5	61.11	90.75
5 Ltd	30	21	4	5	70.00	102.00
5 Bld	28	18	2	8	64.29	95.00
5 Mp	20	12	4	4	60.00	90.25
5 Qyhk	23	15	2	6	65.22	96.00
7 DD	31	26	1	4	83.87	103.25
5 Gwl	25	18	2	6	72.00	100.50
5 JYD	34	24	5	6	70.59	101.75
5 Lgd	36	27	2	7	75.00	99.50
7 Mh	24	19	1	5	79.17	98.75
6 YmM	24	19	2	4	79.17	99.00
5 LGP	42	33	2	8	78.57	108.50
3 CTL	27	13	1	13	48.15	85.50
5 RZD	33	24	3	6	72.73	103.50
5 NHW	40	18	4	18	45.00	83.75
6 LD	26	12	0	14	46.15	84.50

Fauna*	Total taxa	Forest dwellers	Mixed dwellers	Grassland dwellers	Forest dweller (%)	BSC-PPFF
5 SDD	25	12	2	11	48.00	89.50
3 YJG	21	6	2	13	28.57	80.50
5 GLS	34	15	3	16	44.12	86.00
5 SCZ	27	14	1	12	51.85	91.50
3 YS	25	10	1	14	40.00	83.50
5 XGS	33	15	3	15	45.45	90.00
2 SLWS	18	5	1	12	27.78	81.00
5 FYD	27	20	3	4	74.07	100.50
5 LJ	16	8	2	6	50.00	90.00
7 MLS	20	16	2	2	80.00	97.50
5 DC	16	7	2	7	43.75	88.00
5 Hld	11	8	2	1	72.73	92.75
5 Xhs	19	11	2	6	57.89	89.50
5 Zkd U	33	19	2	12	57.58	93.50
5 MHS	21	14	0	7	66.67	97.00
5 Qzas	13	8	1	4	61.54	93.25
5 Ts	17	10	3	4	58.82	94.50
5 Zkd M	21	11	3	7	52.38	92.00
5 Zkd L	36	18	4	14	50.00	89.75
5 Cjw	8	6	1	1	75.00	92.00
5 Dh	18	11	2	5	61.11	90.75
5 Ltd	30	21	4	5	70.00	102.00
5 Bld	28	18	2	8	64.29	95.00
5 Mp	20	12	4	4	60.00	90.25
5 Qyhk	23	15	2	6	65.22	96.00
7 DD	31	26	1	4	83.87	103.25
5 Gwl	25	18	2	6	72.00	100.50
5 JYD	34	24	5	6	70.59	101.75
5 Lgd	36	27	2	7	75.00	99.50
7 Mh	24	19	1	5	79.17	98.75
6 YmM	24	19	2	4	79.17	99.00
5 LGP	42	33	2	8	78.57	108.50
3 CTL	27	13	1	13	48.15	85.50
5 RZD	33	24	3	6	72.73	103.50
5 NHW	40	18	4	18	45.00	83.75
6 LD	26	12	0	14	46.15	84.50

Fauna*	Total taxa	Forest dwellers	Mixed dwellers	Grassland dwellers	Herbivores (%)	Forest dweller (%)	BSC-PPHF
5 SDD	9	3	0	6	36.00	33.33	53.50
3 YJG	13	4	0	9	61.90	30.77	49.50
5 GLS	15	5	0	10	44.12	33.33	50.50
5 SCZ	15	7	0	8	55.56	46.67	55.00
3 YS	19	8	0	11	76.00	42.11	51.00
5 XGS	18	8	0	10	54.55	44.44	54.00
2 SLWS	14	3	1	10	77.78	21.43	46.50
5 FYD	12	8	2	2	44.44	66.67	58.50
5 LJ	12	6	1	5	75.00	50.00	55.00
7 MLS	10	6	2	2	50.00	60.00	56.00
5 DC	11	5	1	5	68.75	45.45	54.50
5 Hld	7	5	1	1	63.64	71.43	56.25
5 Xhs	10	6	1	3	52.63	60.00	54.50
5 Zkd U	13	7	0	6	39.39	53.85	54.50
5 MHS	9	7	0	2	42.86	77.78	60.00
5 Qzas	6	4	0	2	46.15	66.67	56.25
5 Ts	8	5	1	2	47.06	62.50	57.50
5 Zkd M	9	5	0	4	42.86	55.56	56.00
5 Zkd L	13	5	1	7	36.11	38.46	52.25
5 Cjw	4	4	0	0	50.00	100.00	56.50
5 Dh	11	8	1	2	61.11	72.73	56.75
5 Ltd	13	11	1	1	43.33	84.62	62.00
5 Bld	15	8	1	6	53.57	53.33	54.50
5 Mp	10	7	1	2	50.00	70.00	55.75
5 Qyhk	12	8	1	3	52.17	66.67	56.00
7 DD	13	10	0	3	41.94	76.92	60.25
5 Gwl	14	9	1	4	56.00	64.29	59.00
5 JYD	16	12	1	3	47.06	75.00	63.25
5 Lgd	17	12	0	5	47.22	70.59	60.00
7 Mh	15	12	0	3	62.50	80.00	60.25
6 YmM	21	17	1	3	87.50	80.95	63.50
5 LGP	23	17	1	5	54.76	73.91	62.50
3 CTL	15	7	0	8	55.56	46.67	51.50
5 RZD	12	10	0	2	36.36	83.33	60.00
5 NHW	22	9	2	11	55.00	40.91	49.75
6 LD	8	2	0	6	30.77	25.00	50.00