黄土丘陵沟壑区植被自然更新的种源限制因素研究
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摘要
黄土丘陵区是我国水土流失与生态系统退化最为严重的地区之一，也是生态环境建设的重点与难点区域。植被恢复是该区水土流失治理与生态安全的根本途径。但是，该区长期的植被恢复建设效果不佳，人工植被物种单一，保存率低。而植被自然恢复因其适应性与稳定性受到关注，于是提出通过植被自然恢复来控制水土流失，改善生态环境。然而，该区是否具有足够的种源及有效的繁殖体来保障植被自然更新与恢复尚不清楚。所以，本文选择延河流域为研究区域，通过对区域物种库、典型小流域不同立地条件的土壤种子库、幼苗库及营养繁殖体库研究的基础上，从种源、扩散、幼苗建植等方面分析该区植被更新的限制因素，主要研究结果如下：
1.区域物种库具有209种植物，分属于45种群落类型，包含了黄土丘陵沟壑区退耕地植被恢复演替不同阶段的群落类型，如退耕恢复初期的猪毛蒿、赖草群落，进一步发展的长芒草、达乌里胡枝子群落，较高阶段的茭蒿、铁杆蒿、白羊草群落；沟坡残留灌草丛群落，如狼牙刺、虎榛子、丁香、黄刺玫、荆条、小叶锦鸡儿、杠柳、酸枣等群落；乔木林群落，如辽东栎、侧柏、小叶杨群落。这些群落的分布环境包含了峁顶，阴阳梁峁坡、沟坡等生境类型，证明区域物种库能够为研究区不同地形、不同土壤侵蚀程度生境的植被恢复提供种源。
2.土壤种子库特征随立地条件及地上植被类型发生变化，平均密度变化在1188~22560seed/m~2之间，且主要分布于表层土壤中；虽然种子有随土壤侵蚀而流失迁移的现象，但土壤侵蚀没有造成土壤种子库的显著降低。土壤种子库中共有91个物种，退耕地植被恢复演替过程中的主要物种在种子库中分布广泛，并多具有持久土壤种子库，能够为侵蚀干扰环境中的幼苗建植提供保障。
3.幼苗的时空变化主要受到土壤水分有效性的影响，随着雨季水热条件的改善，幼苗密度和物种丰富度均有显著的增加；而且坡面上有利于水分保存的微生境能够增加种子萌发与幼苗存活。幼苗调查中共记录91个物种，其中退耕地植被恢复演替过程中的主要物种在土壤种子库、幼苗库与地上植被中对应出现，说明这些物种能够在自然条件下萌发、建植，并实现植被在侵蚀环境中的自然更新与演替。
4.物种库中有24种多年生禾草、40种多年生草本物种具有营养繁殖能力，而多数乔木、灌木物种具有萌生能力。植被恢复演替过程中的主要营养繁殖物种具有较强的营养繁殖能力，营养繁殖体芽库密度要远远高于对应物种的土壤种子库密度及幼苗密度。营养繁殖物种更能适应存在土壤侵蚀的环境，表现为随着坡度的增加营养繁殖物种在群落中的重要值显著增加，而依靠种子繁殖的物种重要值逐渐降低。
5.在退耕地自然恢复过程中，早期物种具有生产大量易于传播的小种子的特性，能够形成高密度土壤种子库，以幼苗库优势迅速占领生境；演替较高阶段的主要建群种、伴生种亦能生产大量易于传播的种子，具有一定数量的土壤种子库，能够在适宜条件实现幼苗萌发与建植，并且大多主要物种具有营养繁殖能力，有利于在侵蚀干扰条件下利用营养繁殖方式维持并扩展种群。所以，退耕地植被恢复过程中的主要物种不受种源和扩散限制。而沟坡残存灌乔物种，一般零散分布于地势较低的沟坡，再加上其种子重量较大、产量较小，在缺乏有效传播媒介的情况下，长距离传播受到限制。可见，这些沟坡残存灌乔物种在种源和种子扩散上均受到一定的限制。
6.研究区退耕地植被恢复演替过程中的主要草本、半灌木物种基本不受种源与扩散限制，能够在退耕后较短的时间恢复到典型的蒿草群落；但沟坡残存乔、灌木物种由于种源和扩散的限制，其恢复缓慢。因此，在植被恢复建设过程中应加大乡土乔、灌木物种扩散、建植、生长限制因素的研究，以加快该区植被恢复演替进程，改善植被结构，提高群落的水土保持与生态功能。
The hilly-gullied region of the Loess Plateau is well known due to the serious soilerosion and ecosystem degradation. It is also the key region needing much hard work torestore ecological environments. The vegetation is the key factor to control soil erosion andrestore the degraded environments. However, the efficiency of vegetation restoration wasnot entirely satisfactory due to the simple planted species and high mortality rate.Compared with an artificial plant community, a natural vegetation community has a higherpotential for adaptability and stability; thus, natural vegetation rehabilitation has beenproposed to control soil erosion in the Loess Plateau region. But, as an ecological stress,the soil erosion can influence the process of vegetation development includingpost-dispersal movement of seeds, establishment of seedlings, growth and reproduction ofplant. However, in the study region with serious soil erosion, it is still not clear whetherenough species and propagule persist in this field for the vegetation natural recruitment andrestoration. Thus, the regional species pool was studied in the Yanhe watershed. The soilseed bank, seedling bank and the vegetative propagation were studied in a typical smallwatershed with different vegetation and erosion intensities. And then the limiting factor ofthe vegetation recruitment was analyzed on the basis of the species sources, seed dispersaland seedling establishment. The main results as follows:
1. In the regional species pool,209species were recorded and belonged to45types ofcommunities which includes the communities during the different successional stages onabandoned land, such as Artemisia scoparia, Leymus secalinus communities in the earlystage, followed by Stipa bungeana, Lespedeza davurica communities and Artemisiagiraldii, Artemisia gmelinii, Bothriochloa ischcemum communities in the later restorationstage; the remnant communities on the gully slope, such as Sophora davidii, Ostryopsisdavidiana, Syringa oblate, Rosa xanthina, Vitex negundo var. heterophylla, Caragana microphylla, Periploca sepium, Ziziphus jujuba communities; and tree communities, suchas Quercus wutaishanica, Platycladus orientals, Populus simonii communities. Thedistribution habitats of these communities include the hilly top, sunny and shady hillyslope and gully slope. In the regional species pool, the dominant constructive species andcompanion species have a wide distribution and can provide seed sources for naturalrevegetation on the abandoned land with different soil erosion conditions.
2. Characteristics of the soil seed bank changed with the ecological environments andstanding vegetation. The average soil seed bank density varied from1188~22560seed/m2.The soil seed bank mainly distributed in the top soil, but there was no significant seed losswith the soil erosion intensity increasing.91species were recorded in the soil seed bank,and the dominant abandoned successional species distributed widely. Most of the dominantspecies had persistent soil seed bank which can enhance their recolonization in the erodedenvironments.
3. Soil water validity was the main factor influence the seedling changing in spaceand time. The density and species richness of seedling both had significant increase withthe improvement of soil water and temperature in the rain season. On the slope, themicrosite where more soil water can be retained could enhance the seed germination andseedling survival.91species were recorded in the seedling bank and57of these speciescoexisted in the soil seed bank, seedling bank and standing vegetation. This resultsuggested that these species can germinate and establish under the natural conditions, andthe vegetation can be recruited and succession in the eroded environments.
4. In the species pool,24perennial grasses and40perennial herbs have the ability ofvegetative propagation. And many of the wood species have the ability of sprouting. Themain clone species had high density of bud bank. The density of bud bank was higher thanthe density of soil seed bank and seedling bank of the corresponding species. Thevegetative reproduction (versus sexual) is the most effective mechanism for species tocolonize and resist on such erosive degraded areas. The important value of the vegetativereproduction species increased with the slope angle increasing.
5. During the abandoned land restoration, the early successional species always canproduce large number of seeds with small size and form high density soil seed bank whichcan recolonization quickly by the seedling press. The following species also can producelarge number of seeds which dispersal distance and persist in the soil seed bank. As thesame time, most of the dominant species can survival and spread under the harsh habitatsby vegetative reproduction. These results indicated that these species were not limited byseeds sources and dispersal. Due to fragment landform the remnant wood species distributed on the gully slope with low frequency. The seed dispersal faces the stress of soilerosion and gravity when they dispersed from the lower position (gully slope) to higherposition (hilly slope). Furthermore, the wood species always produce big seeds which cannot disperse distance without help. Thus the recolonization of these wood species waslimited by the seeds sources and dispersal.
6. In the study region, the main herb and subshrub species can recolonize theabandoned slope land within decades and form typical steppe community without limit inseeds sources and dispersal. But the woody species recolonize slowly due to the limit ofseeds sources and dispersal. These results suggested that more studies should be done tofind out the limiting factor in seed dispersal, seedling establishment and growth of thenative woody species in order to enhance their restoration on the abandoned slope landwhich can improve the vegetation structure and enhance the ability of control soil erosionand ecological function.
1.区域物种库具有209种植物，分属于45种群落类型，包含了黄土丘陵沟壑区退耕地植被恢复演替不同阶段的群落类型，如退耕恢复初期的猪毛蒿、赖草群落，进一步发展的长芒草、达乌里胡枝子群落，较高阶段的茭蒿、铁杆蒿、白羊草群落；沟坡残留灌草丛群落，如狼牙刺、虎榛子、丁香、黄刺玫、荆条、小叶锦鸡儿、杠柳、酸枣等群落；乔木林群落，如辽东栎、侧柏、小叶杨群落。这些群落的分布环境包含了峁顶，阴阳梁峁坡、沟坡等生境类型，证明区域物种库能够为研究区不同地形、不同土壤侵蚀程度生境的植被恢复提供种源。
2.土壤种子库特征随立地条件及地上植被类型发生变化，平均密度变化在1188~22560seed/m~2之间，且主要分布于表层土壤中；虽然种子有随土壤侵蚀而流失迁移的现象，但土壤侵蚀没有造成土壤种子库的显著降低。土壤种子库中共有91个物种，退耕地植被恢复演替过程中的主要物种在种子库中分布广泛，并多具有持久土壤种子库，能够为侵蚀干扰环境中的幼苗建植提供保障。
3.幼苗的时空变化主要受到土壤水分有效性的影响，随着雨季水热条件的改善，幼苗密度和物种丰富度均有显著的增加；而且坡面上有利于水分保存的微生境能够增加种子萌发与幼苗存活。幼苗调查中共记录91个物种，其中退耕地植被恢复演替过程中的主要物种在土壤种子库、幼苗库与地上植被中对应出现，说明这些物种能够在自然条件下萌发、建植，并实现植被在侵蚀环境中的自然更新与演替。
4.物种库中有24种多年生禾草、40种多年生草本物种具有营养繁殖能力，而多数乔木、灌木物种具有萌生能力。植被恢复演替过程中的主要营养繁殖物种具有较强的营养繁殖能力，营养繁殖体芽库密度要远远高于对应物种的土壤种子库密度及幼苗密度。营养繁殖物种更能适应存在土壤侵蚀的环境，表现为随着坡度的增加营养繁殖物种在群落中的重要值显著增加，而依靠种子繁殖的物种重要值逐渐降低。
5.在退耕地自然恢复过程中，早期物种具有生产大量易于传播的小种子的特性，能够形成高密度土壤种子库，以幼苗库优势迅速占领生境；演替较高阶段的主要建群种、伴生种亦能生产大量易于传播的种子，具有一定数量的土壤种子库，能够在适宜条件实现幼苗萌发与建植，并且大多主要物种具有营养繁殖能力，有利于在侵蚀干扰条件下利用营养繁殖方式维持并扩展种群。所以，退耕地植被恢复过程中的主要物种不受种源和扩散限制。而沟坡残存灌乔物种，一般零散分布于地势较低的沟坡，再加上其种子重量较大、产量较小，在缺乏有效传播媒介的情况下，长距离传播受到限制。可见，这些沟坡残存灌乔物种在种源和种子扩散上均受到一定的限制。
6.研究区退耕地植被恢复演替过程中的主要草本、半灌木物种基本不受种源与扩散限制，能够在退耕后较短的时间恢复到典型的蒿草群落；但沟坡残存乔、灌木物种由于种源和扩散的限制，其恢复缓慢。因此，在植被恢复建设过程中应加大乡土乔、灌木物种扩散、建植、生长限制因素的研究，以加快该区植被恢复演替进程，改善植被结构，提高群落的水土保持与生态功能。
The hilly-gullied region of the Loess Plateau is well known due to the serious soilerosion and ecosystem degradation. It is also the key region needing much hard work torestore ecological environments. The vegetation is the key factor to control soil erosion andrestore the degraded environments. However, the efficiency of vegetation restoration wasnot entirely satisfactory due to the simple planted species and high mortality rate.Compared with an artificial plant community, a natural vegetation community has a higherpotential for adaptability and stability; thus, natural vegetation rehabilitation has beenproposed to control soil erosion in the Loess Plateau region. But, as an ecological stress,the soil erosion can influence the process of vegetation development includingpost-dispersal movement of seeds, establishment of seedlings, growth and reproduction ofplant. However, in the study region with serious soil erosion, it is still not clear whetherenough species and propagule persist in this field for the vegetation natural recruitment andrestoration. Thus, the regional species pool was studied in the Yanhe watershed. The soilseed bank, seedling bank and the vegetative propagation were studied in a typical smallwatershed with different vegetation and erosion intensities. And then the limiting factor ofthe vegetation recruitment was analyzed on the basis of the species sources, seed dispersaland seedling establishment. The main results as follows:
1. In the regional species pool,209species were recorded and belonged to45types ofcommunities which includes the communities during the different successional stages onabandoned land, such as Artemisia scoparia, Leymus secalinus communities in the earlystage, followed by Stipa bungeana, Lespedeza davurica communities and Artemisiagiraldii, Artemisia gmelinii, Bothriochloa ischcemum communities in the later restorationstage; the remnant communities on the gully slope, such as Sophora davidii, Ostryopsisdavidiana, Syringa oblate, Rosa xanthina, Vitex negundo var. heterophylla, Caragana microphylla, Periploca sepium, Ziziphus jujuba communities; and tree communities, suchas Quercus wutaishanica, Platycladus orientals, Populus simonii communities. Thedistribution habitats of these communities include the hilly top, sunny and shady hillyslope and gully slope. In the regional species pool, the dominant constructive species andcompanion species have a wide distribution and can provide seed sources for naturalrevegetation on the abandoned land with different soil erosion conditions.
2. Characteristics of the soil seed bank changed with the ecological environments andstanding vegetation. The average soil seed bank density varied from1188~22560seed/m2.The soil seed bank mainly distributed in the top soil, but there was no significant seed losswith the soil erosion intensity increasing.91species were recorded in the soil seed bank,and the dominant abandoned successional species distributed widely. Most of the dominantspecies had persistent soil seed bank which can enhance their recolonization in the erodedenvironments.
3. Soil water validity was the main factor influence the seedling changing in spaceand time. The density and species richness of seedling both had significant increase withthe improvement of soil water and temperature in the rain season. On the slope, themicrosite where more soil water can be retained could enhance the seed germination andseedling survival.91species were recorded in the seedling bank and57of these speciescoexisted in the soil seed bank, seedling bank and standing vegetation. This resultsuggested that these species can germinate and establish under the natural conditions, andthe vegetation can be recruited and succession in the eroded environments.
4. In the species pool,24perennial grasses and40perennial herbs have the ability ofvegetative propagation. And many of the wood species have the ability of sprouting. Themain clone species had high density of bud bank. The density of bud bank was higher thanthe density of soil seed bank and seedling bank of the corresponding species. Thevegetative reproduction (versus sexual) is the most effective mechanism for species tocolonize and resist on such erosive degraded areas. The important value of the vegetativereproduction species increased with the slope angle increasing.
5. During the abandoned land restoration, the early successional species always canproduce large number of seeds with small size and form high density soil seed bank whichcan recolonization quickly by the seedling press. The following species also can producelarge number of seeds which dispersal distance and persist in the soil seed bank. As thesame time, most of the dominant species can survival and spread under the harsh habitatsby vegetative reproduction. These results indicated that these species were not limited byseeds sources and dispersal. Due to fragment landform the remnant wood species distributed on the gully slope with low frequency. The seed dispersal faces the stress of soilerosion and gravity when they dispersed from the lower position (gully slope) to higherposition (hilly slope). Furthermore, the wood species always produce big seeds which cannot disperse distance without help. Thus the recolonization of these wood species waslimited by the seeds sources and dispersal.
6. In the study region, the main herb and subshrub species can recolonize theabandoned slope land within decades and form typical steppe community without limit inseeds sources and dispersal. But the woody species recolonize slowly due to the limit ofseeds sources and dispersal. These results suggested that more studies should be done tofind out the limiting factor in seed dispersal, seedling establishment and growth of thenative woody species in order to enhance their restoration on the abandoned slope landwhich can improve the vegetation structure and enhance the ability of control soil erosionand ecological function.
引文
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