Characteristics and regeneration potential of soil seed banks in green space of urban parks in Beijing
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摘要:目的
作为潜在的植物群落，土壤种子库在一定程度上代表地上植被的更新演替潜力，研究城市公园绿地土壤种子库特征及其对地上植被和土壤因子的响应，为近自然城市公园的营造提供理论依据。
方法以北京市六环内公园绿地为对象，将其划分为综合公园、郊野公园、文化遗址公园、社区公园4种类型。在随机选取的75个公园内，采用样方法进行地上植被调查，随机法进行土壤采样，土壤种子库采用萌发试验确定，常规方法获取土壤理化指标。
结果（1）北京城市公园土壤种子库共有77种植物，隶属34科74属，以草本植物为主（97.5%），其中菊科物种数最多（16种）。土壤种子库密度在352 ~ 899粒/m2之间，其中菊科种子储量最高，其次为禾本科和豆科，密度最大的物种为马唐（147.11粒/m2），其次为苜蓿（128.40粒/m2）、狗尾草（122.67粒/m2）等。4类城市公园均以一、二年生草本为主，郊野公园的多年生草本相对丰度要高于其他类型公园，其种子库密度显著大于其他类型公园（P < 0.05）。（2）城市公园土壤种子库的Shannon-Wiener多样性指数范围在1.04 ~ 1.46，Simpson优势度指数范围在0.53 ~ 0.62，Patrick丰富度指数范围在4.79 ~ 8.41，Pielou均匀度指数范围在0.74 ~ 0.84，其中郊野公园种子库物种丰富度显著大于其他类型公园（P < 0.05）。（3）相似系数（Sørensen指数）和非度量多维尺度排序分析表明，不同类型公园间土壤种子库、地上植被相似性系数较高，且群落结构差异不显著。4类城市公园土壤种子库与地上植被相似性偏低，相似系数为0.10 ~ 0.18，且土壤种子库与地上植被间群落构成存在明显差异。（4）土壤种子库密度与草本多样性指数、草本物种丰富度指数、土壤铵态氮和非毛管孔隙度呈显著正相关（P < 0.05），与土壤全氮和硝态氮呈显著负相关；土壤种子库物种丰富度指数与土壤全氮和硝态氮呈显著正相关，与草本多样性指数和土壤铵态氮呈显著负相关。土壤种子库均匀度、优势度和多样性指数与灌木物种丰富度指数呈显著正相关，与草本物种丰富度指数呈显著负相关。其中，草本多样性指数和灌木物种丰富度指数对城市公园土壤种子库的密度和物种多样性影响较大。
结论城市公园土壤种子库密度较小，与其对应地上植被间相似性也较低，且以草本植物种子为主；种子库密度和物种多样性主要受地上草本多样性指数和灌木物种丰富度指数影响较大。因此城市公园绿地土壤种子库更新潜力较低，采取人工促进木本植被更新，是营建近自然城市公园的关键。
Abstract:ObjectiveAs a potential plant community, soil seed bank represents the regeneration and succession potential of aboveground vegetation to some extent. To study the characteristics of soil seed banks and their response to aboveground vegetation and soil factors in urban park green spaces can provide theoretical basis for the construction of urban parks near nature.
MethodTaking the green spaces of parks within the Sixth Ring Road of Beijing as the object, they were divided into four types: comprehensive park, country park, cultural heritage park, and community park. In 75 randomly selected parks, aboveground vegetation surveys were carried out by sampling methods and soil sampling was carried out by randomization methods, the soil seed bank was determined by greenhouse germination method, soil physical and chemical indexes were obtained by conventional method.
Result(1) There were 77 species in the soil seed banks of Beijing City Parks, belonging to 34 families and 74 genera. Most of them were herbs (97.5%), with the highest number of species in the Asteraceae (16 species). Density of soil seed bank ranged from 352 to 899 grain/m2, with Asteraceae having the highest seed reserves, followed by Gramineae and Leguminosae. The species with the highest density was Digitaria sanguinalis (147.11 grain/m2), followed by Medicago sativa (128.40 grain/m2), Setaria viridis (122.67 grain/m2) and others. Annal and biennial herbs were dominant in the four types of urban parks. The relative abundance of perennial herbs in country parks was higher than that in other types of parks, and the density of their seed banks was significantly greater than that in other types of parks (P < 0.05). (2) The Shannon-Wiener diversity index of the soil seed banks in urban parks ranged from 1.04 to 1.46, the Simpson dominance index ranged from 0.53 to 0.62, the Patrick richness index ranged from 4.79 to 8.41, and the Pielou evenness index ranged from 0.74 to 0.84, with the species richness of seed banks of country parks significantly greater than that of other types of parks (P < 0.05). (3) Similarity coefficients (Sørensen index) and non-metric multidimensional scaling analysis indicated that the similarity coefficients of soil seed bank and aboveground vegetation among different types of parks were high, and the differences in community structure were not significant. The similarity between soil seed bank and aboveground vegetation in the four types of urban parks was low, with similarity coefficients ranging from 0.10 to 0.18, and there were significant differences in community composition between them. (4) The density of soil seed banks was significantly positively correlated with herb diversity index, herb species richness index, soil ammonium nitrogen, and non-capillary porosity (P < 0.05), and significantly negatively correlated with soil total nitrogen and nitrate nitrogen. The species richness index of soil seed banks was significantly positively correlated with soil total nitrogen and nitrate nitrogen, and significantly negatively correlated with herb diversity index and soil ammonium nitrogen. The soil seed bank evenness, dominance, and diversity indices were significantly positively correlated with shrub species richness index and negatively correlated with herb species richness index. Among them, the herb diversity index and shrub species richness index had a greater impact on the density and species diversity of soil seed bank in urban parks.
ConclusionThe density of soil seed banks in urban parks and the similarity between the soil seed banks and its corresponding aboveground vegetation is low, and the seeds of herb plants are dominated. The density and species diversity of the seed banks are mainly influenced by the diversity index of aboveground herb plants and the richness index of aboveground shrub species. Therefore, the soil seed bank renewal potential of urban parks is low, and artificial promotion of woody vegetation renewal is the key to build a near-natural urban park.
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图 1 北京市六环内公园研究样地分布
Figure 1. Distribution of study sample plots of parks in the Sixth Ring Road of Beijing
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图 2 不同类型城市公园土壤种子库密度
不同小写字母表示不同公园类型间差异显著（P < 0.05）。误差线代表标准误差。下同。Different small letters indicate significant differences among different park types at 0.05 level. The error line represents the standard error. Same as below.
Figure 2. Density of soil seed banks in different types of urban parks
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图 3 不同类型城市公园土壤种子库生活型组成
Figure 3. Life type composition of soil seed banks in different types of urban parks
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图 4 不同类型城市公园土壤种子库物种多样性指标
Figure 4. Species diversity of soil seed banks in different types of urban parks
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图 5 不同类型城市公园土壤种子库与地上植被NMDS排序
Figure 5. NMDS ranking of soil seed banks and aboveground vegetation in different types of urban parks
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图 6 不同类型城市公园土壤种子库与地上植被之间Sørensen相似性系数
Figure 6. Sørensen similarity coefficients between soil seed banks and aboveground vegetation in different types of urban parks
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图 7 土壤种子库特征与环境因子间RDA排序图
Figure 7. RDA ranking diagram between soil seed banks and environmental factors
表 1 公园分类
Table 1 Classification of parks
公园类型
Type of park位置
Location功能特点
Functional characteristics人为管理
Artificial management综合公园
Comprehensive
park中心区
Central zone综合公园一般规模较大；以人工景观为主；具有休闲游憩、生态景观、文化科普、防灾避灾等功能并全面均衡发展；服务半径大
The scale of comprehensive park is generally large, mainly artificial landscape, and has the functions of recreation, ecological landscape, cultural science, disaster prevention and avoidance, and comprehensive and balanced development; the service radius of such park is large管理机制稳定健全，人为干扰强度较大；资源保护措施完善且效果好；绿地养护水平高
The management mechanism is stable and sound, and the intensity of human interference is large; the resource protection measures are perfect and effective; the level of green space maintenance is high文化遗址公园
Cultural
heritage park中心区
Central zone文化遗址公园规模适中；人文与自然景观相结合；具有突出的文化科普和休闲游憩的功能，对非本地游客有重要影响；服务半径适中
The scale of cultural heritage park is moderate, combining humanistic and natural landscapes, with outstanding functions of popularization of culture and recreation, and have an important impact on non-local visitors; the service radius of such park is moderate管理机制稳定健全，人为干扰强度较大；资源保护措施完善且效果好；绿地养护水平高
The management mechanism is stable and sound, and the intensity of human interference is large; the resource protection measures are perfect and effective; the level of green space maintenance is high社区公园
Community park中心区 （居民区附近）
Central zone (near a residential area)社区公园一般规模较小；以人工景观为主；周围居民为主要服务对象，具有休闲放松、生态景观和防灾避灾等功能；服务半径小
The scale of community parks is generally small, mainly artificial landscape, and the surrounding residents as the main service object, with the functions of leisure and relaxation, ecological landscape and disaster prevention and avoidance; the service radius of such park is small管理机制较稳定，人为干扰强度适中；资源保护措施比较完善且效果比较好；绿地养护水平中等
The management mechanism is relatively sound, and the intensity of human interference is moderate; the resource protection measures are sound and the effect is good; the level of green space maintenance is medium郊野公园
Country park近郊
Suburb郊野公园规模较大；以自然景观为主；生态服务功能突出，兼有休憩娱乐、文化科普和防灾减灾等功能；服务半径较大
The scale of country park is large, mainly natural landscape, with outstanding ecological service functions, and functions of leisure and recreation, culture and science popularization, and disaster prevention and mitigation; the service radius of such park is large管理机制有待提升，人为干扰强度较小；资源保护措施有待完善，效果较好；绿地养护水平一般
The management mechanism needs to be improved, and the intensity of human interference is small; the resource protection measures need to be improved and the effect is good; the level of green space maintenance is average注：引自文献[2, 23−25]。Note: cited from reference [2, 23−25]. 
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表 2 不同类型城市公园土壤种子库物种组成和密度
Table 2 Species composition and density of soil seed banks in different types of urban parks
科
Family属
Genus种
Species生活型
Life
form物种密度/（粒·m−2） Species density/(grain·m−2) 综合公园
Comprehensive
park郊野公园
Country
park文化遗址
公园
Cultural
heritage park社区公园
Community
park车前科 Plantaginaceae 车前属 Plantago 车前 Plantago asiatica P 14.81 62.96 14.81 22.22 婆婆纳属 Veronica 蚊母草 Veronica peregrina A 14.81 唇形科 Lamiaceae 薄荷属 Mentha 薄荷 Mentha canadensis P 14.81 111.11 活血丹属 Glechoma 活血丹 Glechoma longituba P 14.81 71.96 14.81 37.04 夏至草属 Lagopsis 夏至草 Lagopsis supina P 29.63 22.22 14.81 14.81 大戟科 Euphorbiaceae 大戟属 Euphorbia 地锦草 Euphorbia humifusa A 29.63 29.63 19.75 铁苋菜属 Acalypha 铁苋菜 Acalypha australis A 22.22 29.63 143.21 22.22 豆科 Fabaceae 胡枝子属 Lespedeza 胡枝子 Lespedeza bicolor S 74.07 决明属 Senna 决明 Senna tora A 103.70 米口袋属 Gueldenstaedtia 少花米口袋 Gueldenstaedtia verna P 59.26 苜蓿属 Medicago 苜蓿 Medicago sativa P 88.89 148.15 禾本科 Poaceae 地毯草属 Axonopus 地毯草 Axonopus compressus P 29.63 狗尾草属 Setaria 狗尾草 Setaria viridis A 18.52 216.67 56.30 122.22 黑麦草属 Lolium 黑麦草 Lolium perenne P 14.81 马唐属 Digitaria 马唐 Digitaria sanguinalis A 287.04 119.75 48.68 140.74 穇属 Eleusine 牛筋草 Eleusine indica A 55.56 129.29 154.07 67.72 早熟禾属 Poa 早熟禾 Poa annua A 79.01 77.04 196.83 29.63 虎耳草科 Saxifragaceae 虎耳草属 Saxifraga 虎耳草 Saxifraga stolonifera P 44.44 29.63 夹竹桃科 Apocynaceae 鹅绒藤属 Cynanchum 萝藦 Cynanchum rostellatum V 14.81 堇菜科 Violaceae 堇菜属 Viola 堇菜 Viola arcuata P 19.75 紫花地丁 Viola philippica P 14.81 14.81 88.89 锦葵科 Grewia 扁担杆属 Grewia 扁担杆 Grewia biloba S 14.81 苘麻属 Abutilon 苘麻 Abutilon theophrasti A 14.81 14.81 田麻属 Corchoropsis 田麻 Corchoropsis crenata A 14.81 菊科 Asteraceae 飞蓬属 Erigeron 小蓬草 Erigeron canadensis A 22.22 一年蓬 Erigeron annuus AB 14.81 鬼针草属 Bidens 鬼针草 Bidens pilosa A 14.81 蒿属 Artemisia 青蒿 Artemisia caruifolia A 59.26 29.63 44.44 黄鹌菜属 Youngia 黄鹌菜 Youngia japonica P 74.07 29.63 蓟属 Cirsium 蓟 Cirsium japonicum P 74.07 29.63 103.70 91.85 假还阳参属 Crepidiastrum 假还阳参 Crepidiastrum lanceolatum P 103.70 59.26 金光菊属 Rudbeckia 黑心金光菊 Rudbeckia hirta AB 14.81 菊三七属 Gynura 白子菜 Gynura divaricata P 59.26 苦苣菜属 Sonchus 长裂苣荬菜 Sonchus brachyotus A 14.81 22.22 苦苣菜 Sonchus oleraceus AB 31.75 47.41 14.81 24.69 苦荬菜属 Ixeris 中华苦荬菜 Ixeris chinensis P 113.58 44.44 鳢肠属 Eclipta 鳢肠 Eclipta prostrata A 14.81 牛膝菊属 Galinsoga 牛膝菊 Galinsoga parviflora A 22.22 29.63 14.81 蒲公英属 Taraxacum 蒲公英 Taraxacum mongolicum P 29.63 29.63 旋覆花属 Inula 旋覆花 Inula japonica P 14.81 苦木科 Simaroubaceae 臭椿属 Ailanthus 臭椿 Ailanthus altissima T 14.81 蓼科 Polygonaceae 千叶兰属 Muehlenbeckia 竹节蓼 Muehlenbeckia platyclada P 44.44 列当科 Orobanchaceae 地黄属 Rehmannia 地黄 Rehmannia glutinosa P 14.81 马齿苋科 Portulacaceae 马齿苋属 Portulaca 马齿苋 Portulaca oleracea A 74.07 48.48 276.54 72.22 木樨科 Oleaceae 梣属 Fraxinus 白蜡树 Fraxinus chinensis T 22.22 29.63 茜草科 Rubiaceae 耳草属 Hedyotis 耳草 Hedyotis auricularia P 14.81 29.63 拉拉藤属 Galium 车轴草 Galium odoratum P 14.81 蔷薇科 Rosaceae 委陵菜属 Potentilla 朝天委陵菜 Potentilla supina AB 207.41 51.03 44.44 37.04 茄科 Solanaceae 茄属 Solanum 龙葵 Solanum nigrum A 14.81 忍冬科 Caprifoliaceae 忍冬属 Lonicera 金银忍冬 Lonicera maackii S 14.81 桑科 Moraceae 构属 Broussonetia 构 Broussonetia papyrifera T 14.81 桑属 Morus 桑 Morus alba T 29.63 14.81 14.81 莎草科 Cyperaceae 莎草属 Cyperus 莎草 Cyperus rotundus P 14.81 水蜈蚣属 Kyllinga 短叶水蜈蚣 Kyllinga brevifolia P 29.63 44.44 14.81 薹草属 Carex 薹草 Carex spp. P 29.63 88.89 十字花科 Brassicaceae 独行菜属 Lepidium 独行菜 Lepidium apetalum AB 29.63 88.89 14.81 37.86 荠属 Capsella 荠 Capsella bursa-pastoris AB 14.81 诸葛菜属 Orychophragmus 诸葛菜 Orychophragmus violaceus AB 29.63 64.20 石竹科 Caryophyllaceae 繁缕属 Stellaria 繁缕 Stellaria media AB 14.81 29.63 488.89 漆姑草属 Sagina 漆姑草 Sagina japonica AB 14.81 天门冬科 Asparagaceae 沿阶草属 Ophiopogon 沿阶草 Ophiopogon bodinieri P 138.27 14.81 55.56 通泉草科 Mazaceae 通泉草属 Mazus 通泉草 Mazus pumilus A 14.81 44.44 14.81 125.93 无患子科 Sapindaceae 栾属 Koelreuteria 栾 Koelreuteria paniculata T 29.63 44.44 22.22 14.81 槭属 Acer 元宝槭 Acer truncatum T 59.26 苋科 Amaranthaceae 沙冰藜属 Bassia 地肤 Bassia scoparia A 74.07 藜属 Chenopodium 藜 Chenopodium album A 29.63 32.59 39.51 48.15 苋属 Amaranthus 凹头苋 Amaranthus blitum A 29.63 44.44 旋花科 Convolvulaceae 打碗花属 Calystegia 打碗花 Calystegia hederacea A 14.81 虎掌藤属 Ipomoea 牵牛 Ipomoea nil A 14.81 荨麻科 Urticaceae 雾水葛属 Pouzolzia 雾水葛 Pouzolzia zeylanica P 14.81 苎麻属 Boehmeria 苎麻 Boehmeria nivea S 29.63 29.63 74.07 44.44 鸭跖草科 Commelinaceae 鸭跖草属 Commelina 鸭跖草 Commelina communis A 14.81 银杏科 Ginkgoaceae 银杏属 Ginkgo 银杏 Ginkgo biloba T 14.81 罂粟科 Papaveraceae 紫堇属 Corydalis 紫堇 Corydalis edulis A 14.81 鸢尾科 Iridaceae 鸢尾属 Iris 鸢尾 Iris tectorum P 29.63 紫草科 Boraginaceae 附地菜属 Trigonotis 附地菜 Trigonotis peduncularis B 107.41 69.63 212.35 80.00 酢浆草科 Oxalidaceae 酢浆草属 Oxalis 酢浆草 Oxalis corniculata P 19.75 37.04 98.77 59.26 注：A.一年生草本；B.二年生草本；AB.一或二年生草本；P.多年生草本；V.藤本；S.灌木；T.乔木。Notes: A, annual herb; B, biennial herb; AB, annual or biennial herb; P, perennial herb; V, vine; S, shrub; T, tree.
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表 3 不同类型城市公园之间土壤种子库、地上植被Sørensen相似性系数
Table 3 Sørensen similarity coefficients of soil seed banks and aboveground vegetation species among different types of urban parks
公园类型 土壤种子库间物种相似性
Species similarity among soil seed banks地上植被间物种相似性
Species similarity among aboveground vegetations综合公园
Comprehensive park郊野公园
Country park文化遗址公园
Cultural heritage park综合公园
Comprehensive park郊野公园
Country park文化遗址公园
Cultural heritage park郊野公园
Country park0.64 0.62 文化遗址公园
Cultural heritage park0.64 0.57 0.60 0.59 社区公园
Community park0.68 0.56 0.57 0.65 0.64 0.60
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