三江平原恢复湿地土壤种子库特征及其与植被的关系

doi:10.3724/SP.J.1258.2012.00763

摘要/Abstract

摘要：

通过幼苗萌发法和样方调查相结合的方法对三江平原不同演替恢复阶段的种子库特征及其与植被的关系进行了研究。将开垦湿地、不同演替恢复阶段湿地以及天然湿地不同土壤层次(0-5、5-10 cm和根茎)的种子库在两种水分条件下(湿润、淹水10 cm)进行萌发处理。结果表明: 随着演替恢复阶段的进行, 种子库的结构和规模逐渐扩大, 地表群落表现出由旱生物种占优势的群落逐渐演变成以小叶章(Calamagrostis angustifolia)占优势的湿生群落的演替趋势。恢复7年湿地、恢复14年湿地、天然湿地土壤种子库萌发物种数分别为24种、29种、39种, 植被物种数为21种、25种、14种。湿地类型、水分条件和土壤层次均显著影响种子库萌发的物种数及幼苗数(p < 0.01)。种子库具有明显的分层现象, 天然湿地0-5 cm土层种子库种子萌发密度是5-10 cm土层的4倍左右, 而恢复湿地仅1.3倍左右, 且土层间萌发物种相似性系数较低。湿润条件下的萌发物种数显著高于淹水条件, 且两种水分条件下萌发物种的生活型不同。由于恢复时间较短, 不同演替恢复阶段的种子库与植被相似性维持在30%以下。湿地中根茎分蘖出大量的湿地物种, 对于小叶章等优势物种的繁殖具有重要作用。研究表明, 在开垦湿地退耕后的次生演替阶段, 种子库能够保持大量的湿地物种, 通过对湿地种子库与植被的关系研究, 能够为三江平原湿地群落演替与湿地恢复提供策略指导。

关键词: 不同演替恢复阶段, 三江平原湿地, 土壤种子库, 植被

Abstract:

Aims Seed banks are an important source of regenerative material in wetlands. Our objective was to study characteristics of the soil seed banks and relationships with the vegetation in different restoration successional stages.
Methods We collected soil seed banks from different depths (0-5 cm, 5-10 cm and rhizomes) from farmed wetland, restored wetland in different successional stages and natural wetland and germinated them under two water regimes (flooded and non-flooded) in a greenhouse. We used quadrats to survey vegetation structure in natural wetland and restored wetlands of different successional stages.
Important findings With increased the restoration time, seed bank composition and size increased, and the vegetation community showed succession from a xerophyte-dominated community to a wetland community dominated by Calamagrostis angustifolia. Number of species germinated from the seed banks was 24, 29 and 39, while 21, 25 and 14 species were recorded in the vegetation of in 7-year restored wetland, 14-year restored wetland and natural wetland, respectively. Wetland type, water regime, depth of burial significantly influenced both the species richness and seedling emergence (p < 0.01). Seed density varied with depth so that seed density was higher in 0-5 cm than in 5-10 cm. The seed density in 0-5 cm was four times as that in 5-10 cm in natural wetland while it was just 1.3 times in restored wetland. The similarity between layers was low. More species germinated under non-flooded condition than flooded and the life forms differed. Because the restoration time was short, the similarity between seed bank and vegetation in different restoration successional stages was always <30%. Many wetland species germinated from rhizome fragments, especially C. angustifolia and other wetland species. Our research suggests that the seed bank could retain and restore several wetland species in different restoration successional stages, which can be valuable for vegetation succession and wetland restoration.

Key words: different restoration succession stages, Sanjiang Plain wetland, soil seed bank, vegetation

王国栋, 吕宪国, 姜明, 王铭, 赵魁义, 娄彦景. 三江平原恢复湿地土壤种子库特征及其与植被的关系. 植物生态学报, 2012, 36(8): 763-773. DOI: 10.3724/SP.J.1258.2012.00763

WANG Guo-Dong, LÜ Xian-Guo, JIANG Ming, WANG Ming, ZHAO Kui-Yi, LOU Yan-Jing. Characteristics of the soil seed banks and relationships with the vegetation in restored wetlands in Sanjiang Plain, northeast of China. Chinese Journal of Plant Ecology, 2012, 36(8): 763-773. DOI: 10.3724/SP.J.1258.2012.00763

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图/表 8

参考文献 33

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生活型 Life form	物种 Species	种子密度 Seed density (individuals·m^-2)
生活型 Life form	物种 Species	开垦湿地 FW	恢复7年湿地 RW-7	恢复14年湿地 RW-14	天然湿地 NW
灌木	细叶沼柳 Salix rosmarinifolia	16 ± 16	144 ± 41 (64)	256 ± 98 (80)	244 ± 49 (40)
Shrub	越桔柳 Salix myrtilloides	0	16 ± 16	32 ± 32	8 ± 8
	粉花绣线菊 Spiraea japonica	0	0	16 ± 16 (16)	96 ± 37 (56)
多年生草本	小叶章 Calamagrostis angustifolia	0	160 ± 40 (16)	608 ± 115 (288)	856 ± 209 (376)
Perennial herb	湿薹草 Carex humida	0	0	0	64 ± 29 (56)
	薹草 Carex dispalata	0	0	96 ± 30 (16)	8 ± 8
	毛薹草 Carex duriuscula	0	0	0	8 ± 8
	球尾花 Lysimachia thyrsiflora	0	64 ± 47	0	0
	五脉山黧豆 Lathyrus quinquenervius	0	0	0	40 ± 27 (40)
	水苏 Stachys japonica	0	64 ± 21 (16)	16 ± 8 (16)	24 ± 12
	三裂慈菇 Sagittaria trifolia	0	16 ± 16	0	0
	龙江凤毛菊 Saussurea amurensis	16 ± 16	0	288 ± 162	40 ± 40 (40)
	宽叶香蒲 Typha latifolia	160 ± 59	176 ± 45	144 ± 49 (64)	200 ± 35 (24)
	苣荬菜 Sonchus brachyotus	0	0	48 ± 32	128 ± 66 (48)
	线叶黑三棱 Sparganium angustifolium	0	0	0	72 ± 29
	菹草 Potamogeton crispus	96 ± 40	192 ± 80	825 ± 360	160 ± 16
	沼泽早熟禾 Poa palustris	0	384 ± 108 (112)	112 ± 80	16 ± 10
	灯芯草 Juncus effusus	0	1712 ± 825 (224)	464 ± 203 (32)	16 ± 10
	地瓜苗 Lycopus lucidus	0	80 ± 29 (64)	32 ± 20 (16)	0
	老鹳草 Geranium wilfordii	0	0	48 ± 48 (48)	0
	泽芹 Sium suave	0	48 ± 48 (48)	0	0
	垂穗粉花地榆 Sanguisorba tenuifolia	0	96 ± 49 (48)	0	16 ± 16 (16)
	山酢浆草 Oxalis acetosella	0	16 ± 16 (16)	0	0
	兴安拉拉藤 Galium dahuricum	0	0	0	8 ± 8
	堇菜 Viola verecunda	0	16 ± 16 (16)	0	8 ± 8
	东北婆婆纳 Veronica rotunda	0	16 ± 16 (16)	0	72 ± 42 (32)
	车前 Plantago asiatica	0	0	0	8 ± 8
	牛鞭草 Hemarthria sibirica	0	0	80 ± 29	24 ± 17
	乌腺金丝桃 Hepericum attenuatum	0	48 ± 32 (16)	16 ± 16	8 ± 8
	苦草 Vallisneria spiralis	0	0	0	8 ± 8
	鳞毛羽节蕨 Gymnocarpium dryopteris	80 ± 16	1216 ± 429 (232)	2896 ± 1374 (50)	5280 ± 2475 (1624)
	还阳参 Crepis crocea	0	0	96 ± 35 (16)	80 ± 37 (32)
	草莓 Fragaria ananassa	0	0	0	8 ± 8
	苋属一种 Amaranthus sp.	0	0	16 ± 16 (16)	0
	大活 Angelica dahurica	0	208 ± 72 (64)	144 ± 73 (32)	112 ± 49 (16)
	金鱼藻 Ceratophyllum demersum	0	0	16 ± 16	216 ± 115
	宽叶山蒿 Artemisia stolonifera	0	0	64 ± 32 (20)	8 ± 8
一、二年生草本	赛繁缕 Stellaria neglecta	0	16 ± 16	80 ± 80	8 ± 8
One biennial herb	东北鼠曲草 Gnaphalium mandshuricum	16 ± 8	176 ± 109	272 ± 94	304 ± 155
	野稗 Echinochloa crusgalli	16 ± 8	0	96 ± 24	104 ± 52
	荠菜 Capsella bursa-pastoris	32 ± 16	352 ± 314	752 ± 497	288 ± 116
	卵穗荸荠 Eleocharis ovata	0	144 ± 98	16 ± 16	16 ± 10
	水蓼 Polygonum hydropiper	0	0	0	24 ± 12
	芒剪股颖 Agrostis trinii	0	0	0	16 ± 16
	灰绿藜 Chenopodium glaucum	0	16 ± 16	48 ± 21	48 ± 25
	狗尾草 Setaria viridis	0	0	48 ± 21	8 ± 8
总数 Total		432 ± 51	5376 ± 1042 (952)	7625 ± 1213 (690)	8632 ± 1526 (2400)

生活型 Life form	物种 Species	种子密度 Seed density (individuals·m^-2)
生活型 Life form	物种 Species	开垦湿地 FW	恢复7年湿地 RW-7	恢复14年湿地 RW-14	天然湿地 NW
灌木	细叶沼柳 Salix rosmarinifolia	16 ± 16	144 ± 41 (64)	256 ± 98 (80)	244 ± 49 (40)
Shrub	越桔柳 Salix myrtilloides	0	16 ± 16	32 ± 32	8 ± 8
	粉花绣线菊 Spiraea japonica	0	0	16 ± 16 (16)	96 ± 37 (56)
多年生草本	小叶章 Calamagrostis angustifolia	0	160 ± 40 (16)	608 ± 115 (288)	856 ± 209 (376)
Perennial herb	湿薹草 Carex humida	0	0	0	64 ± 29 (56)
	薹草 Carex dispalata	0	0	96 ± 30 (16)	8 ± 8
	毛薹草 Carex duriuscula	0	0	0	8 ± 8
	球尾花 Lysimachia thyrsiflora	0	64 ± 47	0	0
	五脉山黧豆 Lathyrus quinquenervius	0	0	0	40 ± 27 (40)
	水苏 Stachys japonica	0	64 ± 21 (16)	16 ± 8 (16)	24 ± 12
	三裂慈菇 Sagittaria trifolia	0	16 ± 16	0	0
	龙江凤毛菊 Saussurea amurensis	16 ± 16	0	288 ± 162	40 ± 40 (40)
	宽叶香蒲 Typha latifolia	160 ± 59	176 ± 45	144 ± 49 (64)	200 ± 35 (24)
	苣荬菜 Sonchus brachyotus	0	0	48 ± 32	128 ± 66 (48)
	线叶黑三棱 Sparganium angustifolium	0	0	0	72 ± 29
	菹草 Potamogeton crispus	96 ± 40	192 ± 80	825 ± 360	160 ± 16
	沼泽早熟禾 Poa palustris	0	384 ± 108 (112)	112 ± 80	16 ± 10
	灯芯草 Juncus effusus	0	1712 ± 825 (224)	464 ± 203 (32)	16 ± 10
	地瓜苗 Lycopus lucidus	0	80 ± 29 (64)	32 ± 20 (16)	0
	老鹳草 Geranium wilfordii	0	0	48 ± 48 (48)	0
	泽芹 Sium suave	0	48 ± 48 (48)	0	0
	垂穗粉花地榆 Sanguisorba tenuifolia	0	96 ± 49 (48)	0	16 ± 16 (16)
	山酢浆草 Oxalis acetosella	0	16 ± 16 (16)	0	0
	兴安拉拉藤 Galium dahuricum	0	0	0	8 ± 8
	堇菜 Viola verecunda	0	16 ± 16 (16)	0	8 ± 8
	东北婆婆纳 Veronica rotunda	0	16 ± 16 (16)	0	72 ± 42 (32)
	车前 Plantago asiatica	0	0	0	8 ± 8
	牛鞭草 Hemarthria sibirica	0	0	80 ± 29	24 ± 17
	乌腺金丝桃 Hepericum attenuatum	0	48 ± 32 (16)	16 ± 16	8 ± 8
	苦草 Vallisneria spiralis	0	0	0	8 ± 8
	鳞毛羽节蕨 Gymnocarpium dryopteris	80 ± 16	1216 ± 429 (232)	2896 ± 1374 (50)	5280 ± 2475 (1624)
	还阳参 Crepis crocea	0	0	96 ± 35 (16)	80 ± 37 (32)
	草莓 Fragaria ananassa	0	0	0	8 ± 8
	苋属一种 Amaranthus sp.	0	0	16 ± 16 (16)	0
	大活 Angelica dahurica	0	208 ± 72 (64)	144 ± 73 (32)	112 ± 49 (16)
	金鱼藻 Ceratophyllum demersum	0	0	16 ± 16	216 ± 115
	宽叶山蒿 Artemisia stolonifera	0	0	64 ± 32 (20)	8 ± 8
一、二年生草本	赛繁缕 Stellaria neglecta	0	16 ± 16	80 ± 80	8 ± 8
One biennial herb	东北鼠曲草 Gnaphalium mandshuricum	16 ± 8	176 ± 109	272 ± 94	304 ± 155
	野稗 Echinochloa crusgalli	16 ± 8	0	96 ± 24	104 ± 52
	荠菜 Capsella bursa-pastoris	32 ± 16	352 ± 314	752 ± 497	288 ± 116
	卵穗荸荠 Eleocharis ovata	0	144 ± 98	16 ± 16	16 ± 10
	水蓼 Polygonum hydropiper	0	0	0	24 ± 12
	芒剪股颖 Agrostis trinii	0	0	0	16 ± 16
	灰绿藜 Chenopodium glaucum	0	16 ± 16	48 ± 21	48 ± 25
	狗尾草 Setaria viridis	0	0	48 ± 21	8 ± 8
总数 Total		432 ± 51	5376 ± 1042 (952)	7625 ± 1213 (690)	8632 ± 1526 (2400)

	物种平均数 Mean number of species		幼苗数 Number of seedlings
	F	p	F	p
湿地类型 Wetland type (wt)	42.880	< 0.001^**	24.138	< 0.001^**
水分条件 Water regime (wr)	481.945	< 0.001^**	246.576	< 0.001^**
土壤层次 Soil layer (sl)	20.770	< 0.001^**	21.920	< 0.001^**
湿地类型×水分条件 wt × wr	32.467	< 0.001^**	13.088	< 0.001^**
湿地类型×土壤层次 wt × sl	1.759	0.111	1.774	0.108
水分条件×土壤层次 wr × sl	1.788	0.171	10.304	< 0.001^**
湿地类型×水分条件×土壤层次 wt × wr × db	1.848	0.093	1.651	0.137

	物种平均数 Mean number of species		幼苗数 Number of seedlings
	F	p	F	p
湿地类型 Wetland type (wt)	42.880	< 0.001^**	24.138	< 0.001^**
水分条件 Water regime (wr)	481.945	< 0.001^**	246.576	< 0.001^**
土壤层次 Soil layer (sl)	20.770	< 0.001^**	21.920	< 0.001^**
湿地类型×水分条件 wt × wr	32.467	< 0.001^**	13.088	< 0.001^**
湿地类型×土壤层次 wt × sl	1.759	0.111	1.774	0.108
水分条件×土壤层次 wr × sl	1.788	0.171	10.304	< 0.001^**
湿地类型×水分条件×土壤层次 wt × wr × db	1.848	0.093	1.651	0.137

物种 Species	湿地类型 Wetland type
物种 Species	天然湿地 NW	恢复7年 RW-7	恢复14年 RW-14
小叶章 Calamagrostis angustifolia^P	1.882^√	0.518^√	1.482^√
宽叶山蒿 Artemisia stolonifer^P	0.173^√	0.672	0.421^√
垂穗粉花地榆 Sanguisorba tenuifolia^P	0.087^√		0.238
芦苇 Phragmites communis^P	0.085	0.239	0.052
粉花绣线菊 Spiraea japonica^P	0.060^√	0.241
地瓜苗 Lycopus lucidus^P	0.042		0.061^√
野火球 Trilolium lupinaster^P	0.039		0.003
赛繁缕 Stellaria neglecta^a	0.036	0.041^√	0.011^√
草莓 Fragaria ananassa^P	0.033^√	0.038	0.057
越桔柳 Salix myrtilloides^P	0.033^√		0.037^√
苣荬菜 Sonchus brachyotus^P	0.028^√	0.032
兴安拉拉藤 Galium dahuricum^P	0.027^√	0.003
沼泽早熟禾 Poa palustris^P	0.02^√	0.215^√	0.172^√
毛薹草 Carex duriuscula^P	0.003^√		0.072
球尾花 Lysimachia thyrsiflora^P		0.007^√	0.003
山杨 Populus davidiana^P		0.152	0.172
毒芹 Cicuta virosa^P		0.007	0.013
千屈菜 Lythrum salicaria^P		0.203	0.132
山酢浆草 Oxalis acetosella^P		0.003^√
细叶沼柳 Salix rosmarinifolia^P		0.083^√	0.152^√
东北鼠曲草 Gnaphalium mandshuricum^a		0.010^√	0.013^√
灰绿黎 Chenopodium glaucum^a		0.003^√	0.006^√
黄莲花 Lysimachia davurica^P		0.007
泽芹 Sium suave^P		0.003^√	0.003
问荆 Equisetum arvense^P		0.054	0.003
狭叶黄岑 Scutellaria regeliana^P		0.008	0.025
五脉山黧豆 Lathyrus quinquenervius^P			0.113
野大豆 Glycine soja^a			0.004
柳蒿 Artemisia integrifolia^P			0.007
老鹳草 Geranium wilfordii^P			0.015^√