Early Miocene flora of central Kazakhstan (Turgai Plateau) and its paleoenvironmental implications

doi:10.1016/j.pld.2019.04.002

Abstract

Abstract: The investigation of the fossil floras from the Turgai plateau (central Kazakhstan) contributes to a better understanding of the origin of the temperate Turgai type flora which spread to Kazakhstan and adjacent areas during the OligoceneeMiocene transition. In this paper, we present the results of a carpological and palynological study of the Kumyrtas flora collected from a flora-bearing horizon of the regional coalbearing Zhilanchik suite, dated to the Aquitanian period. Pollen analysis identified 33 taxa, with are dominated by angiosperms (about 73%) and reflect zonal vegetation. The high percentages of Betula (27%) and Pterocarya (7.5%) that were found in this flora allows comparisons with other Aquitanian floras of Kazakhstan. Based on descriptions of fossil fruits and seeds, we determined that 19 taxa were dominant; these taxa had meosphytic herbaceous components, suggesting mostly edaphic local conditions. The incongruence between the carpological and the pollen records suggests a significant taphonomical effect. Quantitative reconstruction of the palaeoclimate based on pollen records supports-and slightly extends-previous findings based on fossil leaf data, but contradicts findings deduced from the carpological record. Plant Functional Type (PFT) classification was used to characterize the vegetation patterns. Pollen records show that about 45% of diversity relates to the arboreal broadleaved deciduous components and ca. 35% to conifers. Fossil fruit and seed data indicate riverine vegetation with a high diversity of aquatic components and shrub stratum.

Key words: Kazakhstan, Turgai flora, OligoceneeMiocene transition, Palaeoclimate reconstruction, PFT

Popova Svetlana, Utescher Torsten, Averyanova Anna, Tarasevich Valentina, Tropina Polina, Xing Yaowu. Early Miocene flora of central Kazakhstan (Turgai Plateau) and its paleoenvironmental implications[J]. Plant Diversity, 2019, 41(03): 183-197.

Figures/Tables 3

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物种 Species	森林类型 Forest type
物种 Species	苔藓矮林 Dwarf mossy forest	木果柯林 Lithocarpus xylocarpus forest	老龄栎类次生林 Old-aged oak secondary forest	中龄栎类次生林 Middle-aged oak secondary forest	滇山杨次生林 Secondary Populus bonatii forest		旱冬瓜次生林 Secondary Alnus nepalensis forest	云南松次生林 Secondary Pinus yunnanensis forest	厚皮香次生林 Secondary Ternstroemia gymnanthera forest
盖度 Coverage (%)
粉缘绵毛衣 Leioderma sorediatum	0.006 ± 0.003^b	0^a	0^a	0.014 ± 0.005^b	0.011 ± 0.003^b	<0.001^a		<0.001^a		0.059 ± 0.017^c
天蓝猫耳衣 Leptogium azureum	0.017 ± 0.008^ab	0.004 ± 0.002^d	0.009 ± 0.003^b	0.089 ± 0.030^c	0.461 ± 0.096^e	0.125 ± 0.056^abc		0.001 ± 0.001^d		0.046 ± 0.020^ac
网肺衣 Lobaria retigera	0.082 ± 0.039^a	<0.001^b	0.002 ± 0.002^bc	0.063 ± 0.031^a	0.349 ± 0.080^f	0.015 ± 0.010^cde		0.002 ± 0.002^bd		0.042 ± 0.018^ae
双缘牛皮叶 Sticta duplolimbata	0.084 ± 0.034^a	0.003 ± 0.002^b	0.008 ± 0.003^c	0.044 ± 0.009^a	0.052 ± 0.014^a	<0.001^b		0.002 ± 0.002^b		0.025 ± 0.007^ac
频度 Frequency (%)
粉缘绵毛衣 Leioderma sorediatum	0.898 ± 0.519^b	0^a	0^a	2.247 ± 0.678^b	1.580 ± 0.366^b	0.089 ± 0.061^a		0.091 ± 0.062^a		9.162 ± 2.016^c
天蓝猫耳衣 Leptogium azureum	2.221 ± 0.796^a	0.533 ± 0.211^c	2.000 ± 0.605^a	7.941 ± 2.067^b	26.998 ± 2.967^d	10.345 ± 3.773^ab		0.142 ± 0.142^c		6.477 ± 1.975^ab
网肺衣 Lobaria retigera	3.375 ± 1.292^a	0.027 ± 0.027^b	0.200 ± 0.200^bc	3.027 ± 1.036^a	10.352 ± 2.224^e	0.327 ± 0.140^cd		0.142 ± 0.142^bd		2.607 ± 0.878^a
双缘牛皮叶 Sticta duplolimbata	6.949 ± 2.663^ac	0.293 ± 0.193^b	1.800 ± 0.563^d	6.528 ± 0.857^a	8.093 ± 1.762^a	0.089 ± 0.089^b		0.236 ± 0.236^b		3.068 ± 0.895^cd
样地发生率 Occurrence in plots (%)
粉缘绵毛衣 Leioderma sorediatum	50.000	0.000	0.000	60.000	73.333	13.333		13.333		80.000
天蓝猫耳衣 Leptogium azureum	90.000	28.000	90.000	100.000	100.000	66.667		6.667		86.667
网肺衣 Lobaria retigera	70.000	4.000	10.000	80.000	80.000	33.333		6.667		60.000
双缘牛皮叶 Sticta duplolimbata	90.000	16.000	90.000	100.000	86.667	6.667		6.667		60.000
宿主占有率 Occurrence on hosts (%)
粉缘绵毛衣 Leioderma sorediatum	4.000	0.000	0.000	7.556	5.778	0.444		0.444		20.889
天蓝猫耳衣 Leptogium azureum	8.333	2.000	7.667	20.444	55.556	26.222		0.444		12.667
网肺衣 Lobaria retigera	12.333	0.133	1.000	9.778	27.556	1.556		0.444		6.889
双缘牛皮叶 Sticta duplolimbata	20.000	1.200	8.000	20.222	22.889	0.444		0.444		8.222

物种 Species	森林类型 Forest type
物种 Species	苔藓矮林 Dwarf mossy forest	木果柯林 Lithocarpus xylocarpus forest	老龄栎类次生林 Old-aged oak secondary forest	中龄栎类次生林 Middle-aged oak secondary forest	滇山杨次生林 Secondary Populus bonatii forest		旱冬瓜次生林 Secondary Alnus nepalensis forest	云南松次生林 Secondary Pinus yunnanensis forest	厚皮香次生林 Secondary Ternstroemia gymnanthera forest
盖度 Coverage (%)
粉缘绵毛衣 Leioderma sorediatum	0.006 ± 0.003^b	0^a	0^a	0.014 ± 0.005^b	0.011 ± 0.003^b	<0.001^a		<0.001^a		0.059 ± 0.017^c
天蓝猫耳衣 Leptogium azureum	0.017 ± 0.008^ab	0.004 ± 0.002^d	0.009 ± 0.003^b	0.089 ± 0.030^c	0.461 ± 0.096^e	0.125 ± 0.056^abc		0.001 ± 0.001^d		0.046 ± 0.020^ac
网肺衣 Lobaria retigera	0.082 ± 0.039^a	<0.001^b	0.002 ± 0.002^bc	0.063 ± 0.031^a	0.349 ± 0.080^f	0.015 ± 0.010^cde		0.002 ± 0.002^bd		0.042 ± 0.018^ae
双缘牛皮叶 Sticta duplolimbata	0.084 ± 0.034^a	0.003 ± 0.002^b	0.008 ± 0.003^c	0.044 ± 0.009^a	0.052 ± 0.014^a	<0.001^b		0.002 ± 0.002^b		0.025 ± 0.007^ac
频度 Frequency (%)
粉缘绵毛衣 Leioderma sorediatum	0.898 ± 0.519^b	0^a	0^a	2.247 ± 0.678^b	1.580 ± 0.366^b	0.089 ± 0.061^a		0.091 ± 0.062^a		9.162 ± 2.016^c
天蓝猫耳衣 Leptogium azureum	2.221 ± 0.796^a	0.533 ± 0.211^c	2.000 ± 0.605^a	7.941 ± 2.067^b	26.998 ± 2.967^d	10.345 ± 3.773^ab		0.142 ± 0.142^c		6.477 ± 1.975^ab
网肺衣 Lobaria retigera	3.375 ± 1.292^a	0.027 ± 0.027^b	0.200 ± 0.200^bc	3.027 ± 1.036^a	10.352 ± 2.224^e	0.327 ± 0.140^cd		0.142 ± 0.142^bd		2.607 ± 0.878^a
双缘牛皮叶 Sticta duplolimbata	6.949 ± 2.663^ac	0.293 ± 0.193^b	1.800 ± 0.563^d	6.528 ± 0.857^a	8.093 ± 1.762^a	0.089 ± 0.089^b		0.236 ± 0.236^b		3.068 ± 0.895^cd
样地发生率 Occurrence in plots (%)
粉缘绵毛衣 Leioderma sorediatum	50.000	0.000	0.000	60.000	73.333	13.333		13.333		80.000
天蓝猫耳衣 Leptogium azureum	90.000	28.000	90.000	100.000	100.000	66.667		6.667		86.667
网肺衣 Lobaria retigera	70.000	4.000	10.000	80.000	80.000	33.333		6.667		60.000
双缘牛皮叶 Sticta duplolimbata	90.000	16.000	90.000	100.000	86.667	6.667		6.667		60.000
宿主占有率 Occurrence on hosts (%)
粉缘绵毛衣 Leioderma sorediatum	4.000	0.000	0.000	7.556	5.778	0.444		0.444		20.889
天蓝猫耳衣 Leptogium azureum	8.333	2.000	7.667	20.444	55.556	26.222		0.444		12.667
网肺衣 Lobaria retigera	12.333	0.133	1.000	9.778	27.556	1.556		0.444		6.889
双缘牛皮叶 Sticta duplolimbata	20.000	1.200	8.000	20.222	22.889	0.444		0.444		8.222

树种 Tree species	样本数 No. of trees	粉缘绵毛衣 Leioderma sorediatum		天蓝猫耳衣 Leptogium azureum		网肺衣 Lobaria retigera		双缘牛皮叶 Sticta duplolimbata
树种 Tree species	样本数 No. of trees	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)
旱冬瓜 Alnus nepalensis	421	<0.01 (0.17)	0.48	0.27 (4.50)	25.18	0.03 (9.50)	1.43	<0.01 (0.17)	0.48
厚皮香 Ternstroemia gymnanthera	397	0.24 (13.33)	21.91	0.17 (15.33)	11.59	0.18 (18.00)	7.30	0.11 (5.67)	10.08
云南松 Pinus yunnanensis	378	<0.01 (0.33)	0.26	0	0	0	0	0	0
稀有树种 Rare trees	360	0.05 (9.33)	2.78	0.16 (13.17)	13.89	0.08 (8.67)	4.44	0.08 (6.33)	8.06
滇山杨 Populus bonatii	358	0.03 (2.33)	5.03	1.42 (17.67)	68.72	1.24 (52.00)	35.20	0.16 (3.67)	27.65
硬壳柯 Lithocarpus hancei	289	0.04 (2.17)	7.27	0.28 (6.50)	23.88	0.11 (9.67)	7.61	0.16 (10.00)	24.57
云南越桔 Vaccinium duclouxii	280	0.01 (1.67)	1.07	0	0	0.01 (4.00)	0.36	<0.01 (0.33)	1.07
珊瑚冬青 Ilex corallina	106	0.01 (0.67)	3.77	0.06 (2.00)	7.55	0	0	0.03 (0.67)	6.60
珍珠花 Lyonia ovalifolia	96	0.03 (0.67)	8.33	0.06 (1.33)	7.29	<0.01 (0.33)	1.04	0.02 (1.17)	6.25
变色锥 Castanopsis wattii	95	0	0	0.01 (0.50)	2.11	0.01 (0.50)	1.05	0.01 (0.67)	1.05
露珠杜鹃 Rhododendron irroratum	92	0.02 (1.00)	4.35	0.01 (1.00)	1.09	0.08 (2.33)	9.78	0.16 (3.00)	17.39
硬叶柯 Lithocarpus crassifolius	82	0.02 (1.33)	3.66	0.06 (1.17)	12.20	0.58 (21.00)	17.07	0.18 (2.17)	21.95
南洋木荷 Schima noronhae	76	0	0	0.02 (0.67)	2.63	0	0	<0.01 (0.33)	1.32
翅柄紫茎 Stewartia pteropetiolata	75	0	0	0	0	0.02 (1.33)	1.33	<0.01 (0.17)	1.33
木果柯 Lithocarpus xylocarpus	70	0	0	<0.01 (0.67)	1.43	<0.01 (0.33)	1.43	<0.01 (0.33)	1.43
薄叶马银花 Rhododendron leptothrium	69	0	0	0.08 (2.33)	8.70	0.02 (1.00)	2.90	0.07 (3.67)	5.80
南亚枇杷 Eriobotrya bengalensis	56	0.01 (0.67)	1.79	0.02 (0.67)	3.57	0.04 (2.00)	3.57	0.01 (0.17)	3.57
滇润楠 Machilus yunnanensis	48	0	0	0.05 (0.83)	14.58	0	0	0.03 (0.67)	8.33
景东冬青 Ilex gintungensis	46	0	0	<0.01 (0.17)	2.17	0	0	0	0
麻子壳柯 Lithocarpus variolosus	46	0.02 (0.17)	10.87	0.08 (0.83)	23.91	0.16 (2.17)	23.91	0.09 (0.83)	23.91
山矾 Symplocos sumuntia	45	0	0	0.04 (0.83)	6.67	0	0	0.01 (0.50)	2.22
红花木莲 Manglietia insignis	42	0	0	<0.01 (0.17)	2.38	0	0	0.01 (0.33)	2.38
云南桤叶树 Clethra delavayi	37	0.04 (0.67)	8.11	0.03 (0.33)	8.11	0.24 (4.67)	18.92	0.67 (6.67)	27.03
小花山茶 Camellia forrestii	36	0	0	0	0	0	0	0	0

树种 Tree species	样本数 No. of trees	粉缘绵毛衣 Leioderma sorediatum		天蓝猫耳衣 Leptogium azureum		网肺衣 Lobaria retigera		双缘牛皮叶 Sticta duplolimbata
树种 Tree species	样本数 No. of trees	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)	盖度 Coverage (%)	宿主占有率 Occurrence on hosts (%)
旱冬瓜 Alnus nepalensis	421	<0.01 (0.17)	0.48	0.27 (4.50)	25.18	0.03 (9.50)	1.43	<0.01 (0.17)	0.48
厚皮香 Ternstroemia gymnanthera	397	0.24 (13.33)	21.91	0.17 (15.33)	11.59	0.18 (18.00)	7.30	0.11 (5.67)	10.08
云南松 Pinus yunnanensis	378	<0.01 (0.33)	0.26	0	0	0	0	0	0
稀有树种 Rare trees	360	0.05 (9.33)	2.78	0.16 (13.17)	13.89	0.08 (8.67)	4.44	0.08 (6.33)	8.06
滇山杨 Populus bonatii	358	0.03 (2.33)	5.03	1.42 (17.67)	68.72	1.24 (52.00)	35.20	0.16 (3.67)	27.65
硬壳柯 Lithocarpus hancei	289	0.04 (2.17)	7.27	0.28 (6.50)	23.88	0.11 (9.67)	7.61	0.16 (10.00)	24.57
云南越桔 Vaccinium duclouxii	280	0.01 (1.67)	1.07	0	0	0.01 (4.00)	0.36	<0.01 (0.33)	1.07
珊瑚冬青 Ilex corallina	106	0.01 (0.67)	3.77	0.06 (2.00)	7.55	0	0	0.03 (0.67)	6.60
珍珠花 Lyonia ovalifolia	96	0.03 (0.67)	8.33	0.06 (1.33)	7.29	<0.01 (0.33)	1.04	0.02 (1.17)	6.25
变色锥 Castanopsis wattii	95	0	0	0.01 (0.50)	2.11	0.01 (0.50)	1.05	0.01 (0.67)	1.05
露珠杜鹃 Rhododendron irroratum	92	0.02 (1.00)	4.35	0.01 (1.00)	1.09	0.08 (2.33)	9.78	0.16 (3.00)	17.39
硬叶柯 Lithocarpus crassifolius	82	0.02 (1.33)	3.66	0.06 (1.17)	12.20	0.58 (21.00)	17.07	0.18 (2.17)	21.95
南洋木荷 Schima noronhae	76	0	0	0.02 (0.67)	2.63	0	0	<0.01 (0.33)	1.32
翅柄紫茎 Stewartia pteropetiolata	75	0	0	0	0	0.02 (1.33)	1.33	<0.01 (0.17)	1.33
木果柯 Lithocarpus xylocarpus	70	0	0	<0.01 (0.67)	1.43	<0.01 (0.33)	1.43	<0.01 (0.33)	1.43
薄叶马银花 Rhododendron leptothrium	69	0	0	0.08 (2.33)	8.70	0.02 (1.00)	2.90	0.07 (3.67)	5.80
南亚枇杷 Eriobotrya bengalensis	56	0.01 (0.67)	1.79	0.02 (0.67)	3.57	0.04 (2.00)	3.57	0.01 (0.17)	3.57
滇润楠 Machilus yunnanensis	48	0	0	0.05 (0.83)	14.58	0	0	0.03 (0.67)	8.33
景东冬青 Ilex gintungensis	46	0	0	<0.01 (0.17)	2.17	0	0	0	0
麻子壳柯 Lithocarpus variolosus	46	0.02 (0.17)	10.87	0.08 (0.83)	23.91	0.16 (2.17)	23.91	0.09 (0.83)	23.91
山矾 Symplocos sumuntia	45	0	0	0.04 (0.83)	6.67	0	0	0.01 (0.50)	2.22
红花木莲 Manglietia insignis	42	0	0	<0.01 (0.17)	2.38	0	0	0.01 (0.33)	2.38
云南桤叶树 Clethra delavayi	37	0.04 (0.67)	8.11	0.03 (0.33)	8.11	0.24 (4.67)	18.92	0.67 (6.67)	27.03
小花山茶 Camellia forrestii	36	0	0	0	0	0	0	0	0

地衣物种 Lichen species	参数 Variable	林龄 Stand age	林冠开阔度 Canopy openness	基面积 Basal area	胸径 DBH	最大胸径 Max DBH	树木密度 Tree density	树种丰富度 Tree richness	树种辛普森指数 Tree Simpson index
线性回归分析 Single linear regression
粉缘绵毛衣 Leioderma sorediatum	盖度 Coverage	0.05^{**}		0.03^*	0.16^{***}	0.09^{***}	0.28^{***}
粉缘绵毛衣 Leioderma sorediatum	频度 Frequency	0.07^{**}		0.03^*	0.20^{***}	0.11^{***}	0.44^{***}
天蓝猫耳衣 Leptogium azureum	盖度 Coverage	0.07^{**}	0.03^*	0.06^{**}		0.05^*
天蓝猫耳衣 Leptogium azureum	频度 Frequency	0.12^{***}	0.05^{**}	0.09^{***}	0.03^*	0.07^{**}
网肺衣 Lobaria retigera	盖度 Coverage	0.03^*	0.07^{**}	0.07^{**}	0.06^{**}	0.05^{**}
网肺衣 Lobaria retigera	频度 Frequency	0.04^*	0.03^*	0.04^*	0.08^{***}	0.07^{**}
双缘牛皮叶 Sticta duplolimbata	盖度 Coverage				0.08^{**}	0.04^*		0.03^*	0.03_*
双缘牛皮叶 Sticta duplolimbata	频度 Frequency				0.10^{***}	0.06^{**}		0.03^*	0.06^{**}
多项式回归分析 Polynomial regression
粉缘绵毛衣 Leioderma sorediatum	盖度 Coverage	0.12^{***}	0.13^{***}		0.31^{***}	0.14^{***}	0.30^{***}		0.04^*
粉缘绵毛衣 Leioderma sorediatum	频度 Frequency	0.16^{***}	0.18^{***}		0.40^{***}	0.18^{***}	0.43^{***}		0.08^{**}
天蓝猫耳衣 Leptogium azureum	盖度 Coverage	0.07^{**}	0.05^*	0.07^{**}		0.04^*
天蓝猫耳衣 Leptogium azureum	频度 Frequency	0.12^{***}	0.13^{***}	0.09^{**}		0.07^{**}
网肺衣 Lobaria retigera	盖度 Coverage	0.04^*	0.07^{**}	0.08^{**}	0.05^*	0.05^*
网肺衣 Lobaria retigera	频度 Frequency	0.04^*	0.12^{***}	0.03^*	0.07^{**}	0.08^{**}	0.05^*		0.04^*
双缘牛皮叶 Sticta duplolimbata	盖度 Coverage		0.20^{***}		0.08^{**}	0.04^*	0.12^{***}	0.04^*	0.04^*
双缘牛皮叶 Sticta duplolimbata	频度 Frequency		0.25^{***}		0.09^{**}	0.05^*	0.14^{***}	0.05^{**}	0.07^{**}