植物种群更新的补充限制

doi:10.17521/cjpe.2020.0246

摘要/Abstract

摘要：

补充限制基于生态位理论, 从种子萌发、幼苗存活和生长、繁殖体扩散等生活史阶段的种群统计特征及环境因素(土壤水分、养分、凋落物等)着手, 探讨种群的更新问题。种源限制和微生境限制是补充限制理论研究的核心内容, 但是哪个更为重要并没有统一的结论。种源限制与种子生产、土壤种子库和地下芽库中的繁殖体数量不足有关。其中, 气候的年际波动、土壤种子库寿命和动物的捕食都会影响种子生产在种群更新中的作用; 土壤种子库常被视为种群更新的保险库, 与地上种子雨共同促进种群更新, 但是, 如果土壤里种子具有较高的死亡率和休眠率, 将会降低种子库的作用; 地下芽库及其产生的无性分株对于种群更新的意义更多地体现在干扰后种群更强的恢复能力上。扩散限制是种群更新中的普遍现象, 与种子产量、散布能力、传播媒介、幼苗密度等因素有关。微生境限制主要表现为水分、养分、凋落物等非生物因素以及竞争、捕食等生物因素对种子的活力、萌发性、幼苗的存活力、物质分配等过程的影响, 其重要性随着植物生活史阶段而发生变化。未来需要进行综合的、长期的实验, 并应着重加强种源限制及相关生态过程的进化与生态相结合的机理性研究, 从而更深刻地认识和理解种群更新问题, 建立更为综合、系统的种群更新理论体系。

关键词: 种源限制, 微生境限制, 幼苗存活, 地下芽库, 扩散限制

Abstract:

Based on niche theory, recruitment limitation is to discuss population regeneration through the population demographic characteristics in different life stages, such as seed germination, seedling survival and growth, propagule dispersal, and varied environmental factors (soil moisture, nutrients, litter, etc.). The core of recruitment limitation consists of seed limitation and microsite limitation; however, there is no universal conclusion on which of them is more dominant. Seed limitation is related to insufficient propagules of seed production, soil seed bank and underground bud bank. Specifically speaking, inter-annual climate fluctuation, soil seed bank life time and animal predation can lead to seed limitation. Soil seed bank is often regarded as the vault of population recruitment, which promotes population recruitment together with the above ground seed rain. In contrast, if soil seeds have high death rate and dormancy rate, the role of seed bank will decline. The contribution of underground bud bank and its clonal ramets for population regeneration are more reflected on stronger resilience to disturbance. Dispersal limitation is a common phenomenon in population regeneration, which is related to many factors including seed production, dispersal ability, spreading media and seedling density. In terms of microsite limitation, embodying in how abiotic factors such as water, nutrients, litters, and biotic factors such as competition and predation effect on seed vigor, germination, seedling viability, and biomass allocation, its importance often varies with the stages of plant life history. In the future, it is necessary to conduct comprehensive and long-term experiments to focus on the underlying mechanisms involving evolution and ecology on seed limitation and its associated ecological processes, so as to obtain a more profound understanding of population recruitment, and establish a more comprehensive and systematic framework of population recruitment.

Key words: seed limitation, microsite limitation, seedling survival, belowground bud-bank, dispersal limitation

韩大勇, 张维, 努尔买买提•依力亚斯, 杨允菲. 植物种群更新的补充限制. 植物生态学报, 2021, 45(1): 1-12. DOI: 10.17521/cjpe.2020.0246

HAN Da-Yong, ZHANG Wei, Nuermaimaiti YILIYASI, YANG Yun-Fei. Recruitment limitation of plant population regeneration. Chinese Journal of Plant Ecology, 2021, 45(1): 1-12. DOI: 10.17521/cjpe.2020.0246

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0246

https://www.plant-ecology.com/CN/Y2021/V45/I1/1

图/表 1

参考文献 110

[1]	Ascoli D, Vacchiano G, Maringer J, Bovio G, Conedera M (2015). The synchronicity of masting and intermediate severity fire effects favors beech recruitment. Forest Ecology and Management, 353,126-135. DOI URL
[2]	Barrett JP, Silander Jr JA (1992). Seedling recruitment limitation in white clover (Trifolium repens; Leguminosae). American Journal of Botany, 79,643-649. DOI URL
[3]	Benson EJ, Hartnett DC, Mann HK (2004). Belowground bud banks and meristem limitation in tallgrass prairie plant populations. American Journal of Botany, 91,416-421. DOI URL
[4]	Bogdziewicz M, Espelta JM, Muñoz A, Aparicio JM, Bonal R (2018). Effectiveness of predator satiation in masting oaks is negatively affected by conspecific density. Oecologia, 186,983-993. DOI PMID
[5]	Callaway RM (1992). Effect of shrubs on recruitment of Quercus douglasii and Quercus lobata in California. Ecology, 73,2118-2128. DOI URL
[6]	Calviño-Cancela M (2007). Seed and microsite limitations of recruitment and the impacts of post-dispersal seed predation at the within population level. Plant Ecology, 192,35-44. DOI URL
[7]	Cárate-Tandalla D, Leuschner C, Homeier J (2015). Performance of seedlings of a shade-tolerant tropical tree species after moderate addition of N and P. Frontiers in Earth Science, 3,75. DOI: 10.3389/feart.2015.00075. DOI
[8]	Caughlin TT, Ferguson JM, Lichstein JW, Zuidema PA, Bunyavejchewin S, Levey DJ (2015). Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages. Proceedings of the Royal Society B: Biological Sciences, 282,20142095. DOI: 10.1098/rspb.2014.2095. DOI URL
[9]	Cheplick GP (1992). Sibling competition in plants. Journal of Ecology, 80,567-575. DOI URL
[10]	Clark CJ, Poulsen JR, Levey DJ (2013). Roles of seed and establishment limitation in determining patterns of afrotropical tree recruitment. PLOS ONE, 8,e63330. DOI: 10.1371/journal.pone.0063330. DOI URL
[11]	Clark CJ, Poulsen JR, Levey DJ, Osenberg CW (2007). Are plant populations seed limited? A critique and meta-analysis of seed addition experiments. The American Naturalist, 170,128-142. PMID
[12]	Clark JS, Silman M, Kern R, Macklin E, HilleRisLambers J (1999). Seed dispersal near and far: patterns across temperate and tropical forests. Ecology, 80,1475-1494. DOI URL
[13]	Clements H, Bierzychudek P (2017). Can the persistent seed bank contribute to the passive restoration of urban forest fragments after invasive species removal? Ecological Restoration, 35,156-166. DOI URL
[14]	Coelho FF, Capelo C, Figueira JEC (2008). Seedlings and ramets recruitment in two rhizomatous species of Rupestrian grasslands: Leiothrix curvifolia var. lanuginosa and Leiothrix crassifolia (Eriocaulaceae). Flora-Morphology, Distribution, Functional Ecology of Plants, 203,152-161.
[15]	Connell JH (1971). On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees//Den Boer PJ, Gradwell GR. Dynamics of Populations. Centre for Agricultural Publishing and Documentation, Wageningen, the Netherlands. 298-312.
[16]	Crawley MJ (1990). The population dynamics of plants. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences, 330,125-140.
[17]	Cui W, Zhao LP, Zhao FR (2017). Effects of fencing and grazing management on bud bank in a semiarid steppe on the Loess Plateau. Pratacultural Science, 34,9-15.
	[ 崔伟, 赵凌平, 赵芙蓉 (2017). 封育和放牧对黄土高原典型草原芽库的影响. 草业科学, 34,9-15.]
[18]	Dalgleish HJ, Hartnett DC (2009). The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography. Plant Ecology, 201,411-420. DOI URL
[19]	Dalgleish HJ, Ott JP, Setshogo MP, Hartnett DC (2012). Inter- specific variation in bud banks and flowering effort among semi-arid African savanna grasses. South African Journal of Botany, 83,127-133. DOI URL
[20]	Dalling JW, Muller-Landau HC, Wright SJ, Hubbell SP (2002). Role of dispersal in the recruitment limitation of neotropical pioneer species. Journal of Ecology, 90,714-727. DOI URL
[21]	de la Peña-Domene M, Martínez-Garza C (2018). Integrating density into dispersal and establishment limitation equations in tropical forests. Forests, 9,570. DOI: 10.3390/ f9090570. DOI URL
[22]	Ding Y, Zang RG, Letcher SG, Liu WD, Lu XH (2016). Aboveground and belowground competition affect seedling performance and allometry in a tropical monsoon forest. New Forests, 47,529-540. DOI URL
[23]	Duclos V, Boudreau S, Chapman CA (2013). Shrub cover influence on seedling growth and survival following logging of a tropical forest. Biotropica, 45,419-426. DOI URL
[24]	Ehrlén J, Eriksson O (2000). Dispersal limitation and patch occupancy in forest herbs. Ecology, 81,1667-1674. DOI URL
[25]	Ehrlén J, Münzbergová Z, Diekmann M, Eriksson O (2006). Long-term assessment of seed limitation in plants: results from an 11-year experiment. Journal of Ecology, 94,1224-1232. DOI URL
[26]	Eriksson O, Ehrlén J (1992). Seed and microsite limitation of recruitment in plant populations. Oecologia, 91,360-364. DOI PMID
[27]	Eskelinen A, Virtanen R (2005). Local and regional processes in low-productive mountain plant communities: the roles of seed and microsite limitation in relation to grazing. Oikos, 110,360-368. DOI URL
[28]	Fischer C, Kollmann J, Wagner TC (2015). How does the seed fate of Crotalaria podocarpa DC, a highly competitive herbaceous legume in arid rangelands, contribute to its establishment probability? Perspectives in Plant Ecology, Evolution and Systematics, 17,405-411. DOI URL
[29]	Fraaije RGA, ter Braak CJF, Verduyn B, Verhoeven JTA, Soons MB (2015). Dispersal versus environmental filtering in a dynamic system: drivers of vegetation patterns and diversity along stream riparian gradients. Journal of Ecology, 103,1634-1646. DOI URL
[30]	García-Fayos P, Gasque M (2006). Seed vs. microsite limitation for seedling emergence in the perennial grass Stipa tenacissima L. (Poaceae). Acta Oecologica, 30,276-282. DOI URL
[31]	García-Meza D, Martorell C (2016). A trade off in the absolute magnitudes of seed and microsite limitations and their effects on population regulation. Oikos, 125,308-314. DOI URL
[32]	Gavinet J, Prévosto B, Fernandez C (2016). Do shrubs facilitate oak seedling establishment in Mediterranean pine forest understory? Forest Ecology and Management, 381,289-296. DOI URL
[33]	Greig-Smith J, Sagar GR (1981). Biological causes of rarity in Carlina vulgaris//Synge H. Biological Aspects of Rare Plant Conservation. Wiley, Chichester.389-400.
[34]	Gu C, Zhao TQ, Wang YT, Wang X, Jia LX, Zhao ML (2017). The response of growth and reproduction for Stipa breviflora to different stocking rates. Ecology and Environmental Sciences, 26,36-42.
	[ 古琛, 赵天启, 王亚婷, 王玺, 贾丽欣, 赵萌莉 (2017). 短花针茅生长和繁殖策略对载畜率的响应. 生态环境学报, 26,36-42.]
[35]	Guo LJ, Cao HW, Xu WH, Tian YQ, Xiao ZP (2017). Seed rain, soil seed bank and quantitative dynamics of seedlings of Emmenopterys henryi populations in different altitude regions. Bulletin of Botanical Research, 37,377-386.
	[ 郭连金, 曹昊玮, 徐卫红, 田玉清, 肖志鹏 (2017). 香果树(Emmenopterys henryi)种群种子雨、种子库及实生苗数量的海拔梯度变化. 植物研究, 37,377-386.]
[36]	Halvorson JJ, Belesky DP, West MS (2017). Inhibition of forage seed germination by leaf litter extracts of overstory hardwoods used in silvopastoral systems. Agroforestry Systems, 91,69-83. DOI URL
[37]	Han DY, Zhao Y, Ahan J, Zhang W, Yang YF (2019). Age structure of Caragana aurantiaca young populations in the Yili valley, Xinjiang. Chinese Journal of Ecology, 38,3601-3608.
	[ 韩大勇, 赵玉, 贾娜尔•阿汗, 张维, 杨允菲 (2019). 新疆伊犁河谷镰叶锦鸡儿(Caragana aurantiaca)幼株种群年龄结构. 生态学杂志, 38,3601-3608.]
[38]	Hartnett DC, Setshogo MP, Dalgleish HJ (2006). Bud banks of perennial savanna grasses in Botswana. African Journal of Ecology, 44,256-263. DOI URL
[39]	Hawthorn WR, Cavers PB (1976). Population dynamics of the perennial herbs Plantago major L. and P. rugelii Decne. The Journal of Ecology, 64,511-527. DOI URL
[40]	Hou ZZ, He S, Wang FL, Zhang AQ (2013). Dynamics of rare clonal plant Linnaea borealis L. sexual reproduction and related affecting factors. Chinese Journal of Ecology, 32,3167-3172.
	[ 侯真珍, 何爽, 王风雷, 张爱勤 (2013). 珍稀克隆植物北极花有性繁殖动态及影响因素. 生态学杂志, 32,3167-3172.]
[41]	Hu JJ, Luo CC, Turkington R, Zhou ZK (2016). Effects of herbivores and litter on Lithocarpus hancei seed germination and seedling survival in the understorey of a high diversity forest in SW China. Plant Ecology, 217,1429-1440. DOI URL
[42]	Ikegami M, Whigham DF, Werger MJA (2012). Effects of local density of clonal plants on their sexual and vegetative propagation strategies in a lattice structure model. Ecological Modelling, 234,51-59. DOI URL
[43]	Janzen DH (1970). Herbivores and the number of tree species in tropical forests. The American Naturalist, 104,501-528. DOI URL
[44]	Jiao DZ, Yao L, Huang ZY, Yang YF (2015). Bud population dynamics of Phragmites australis in heterogeneous habitats of Northeast grassland, China. Chinese Journal of Applied Ecology, 26,404-410.
	[ 焦德志, 么璐, 黄曌月, 杨允菲 (2015). 东北草地异质生境芦苇芽种群动态. 应用生态学报, 26,404-410.]
[45]	Jiménez-Alfaro B, Silveira FAO, Fidelis A, Poschlod P, Commander LE (2016). Seed germination traits can contribute better to plant community ecology. Journal of Vegetation Science, 27,637-645. DOI URL
[46]	Joubert DF, Smit GN, Hoffman MT (2013). The influence of rainfall, competition and predation on seed production, germination and establishment of an encroaching Acacia in an arid Namibian savanna. Journal of Arid Environments, 91,7-13. DOI URL
[47]	Kalle H, Ari-Pekka H, Pasi R, Juha T (2009). Seed introduction and gap creation facilitate restoration of meadow species richness. Journal for Nature Conservation, 17,236-244. DOI URL
[48]	Koyama H (1998). Seed heteromorphism and timing of germination, their mechanisms and adaptive significance. Japanese Journal of Ecology, 48,129-142.
[49]	Kupferschmid AD, Stampfli A, Newbery DM (2000). Dispersal and microsite limitation in an abandoned calcareous grassland of the southern prealps. Folia Geobotanica, 35,125-141. DOI URL
[50]	la Mantia T, Rühl J, Massa B, Pipitone S, Lo Verde G, Bueno RS (2019). Vertebrate-mediated seed rain and artificial perches contribute to overcome seed dispersal limitation in a Mediterranean old field. Restoration Ecology, 27,1393-1400. DOI URL
[51]	Larson JE, Funk JL (2016). Regeneration: an overlooked aspect of trait-based plant community assembly models. Journal of Ecology, 104,1284-1298. DOI URL
[52]	Lee EH, Lee BE, Kim JG (2018). Effects of water levels and soil nutrients on the growth of Iris laevigata seedlings. Journal of Ecology and Environment, 42,5. DOI: 10.1186/s41610-018-0065-4. DOI URL
[53]	Li AD, Peng X, Zhang JL (2014). Study on plant regeneration bank in burned land after forest fire in karst areas. Guizhou Science, 32,40-46.
	[ 李安定, 彭熙, 张建利 (2014). 喀斯特山区森林火灾后火烧迹地植物更新库研究. 贵州科学, 32,40-46.]
[54]	Li HY, Yang YF (2011). Age structures of Spodiopogon sibiricus populations on weedy meadow and elm woodland in Songnen Plain, Northeast China. Chinese Journal of Applied Ecology, 22,1982-1986.
	[ 李海燕, 杨允菲 (2011). 松嫩平原杂类草草甸和榆树疏林草原大油芒种群的年龄结构. 应用生态学报, 22,1982-1986.]
[55]	Liu MH, Ye J, Wen B (2016). Seed germination and seedling survival of Baccaurea ramiflora and Saprosma ternata in Xishuangbanna tropical rainforests under habitat heterogeneity. Acta Botanica Boreali-Occidentalia Sinica, 36,1654-1661.
	[ 刘明航, 叶娟, 文彬 (2016). 热带雨林林内小生境对木奶果和染木种子萌发与幼苗存活的影响. 西北植物学报, 36,1654-1661.]
[56]	Lu JM, Johnson DJ, Qiao XJ, Lu ZJ, Wang QG, Jiang MX (2015). Density dependence and habitat preference shape seedling survival in a subtropical forest in central China. Journal of Plant Ecology, 8,568-577.
[57]	Lucas-Borja ME, Candel-Pérez D, García Morote FA, Onkelinx T, Tíscar PA, Balandier P (2016). Pinus nigra Arn. ssp. salzmannii seedling recruitment is affected by stand basal area, shrub cover and climate interactions. Annals of Forest Science, 73,649-656. DOI URL
[58]	Luo Y, He F, Yu S (2013). Recruitment limitation of dominant tree species with varying seed masses in a subtropical evergreen broad-leaved forest. Community Ecology, 14,189-195. DOI URL
[59]	Ma DX, Liu RL (2012). Comparative study on germination of Bretschneidara sinensis seeds treated with solution extracted from forest litter and other methods. Forest Research, 25,632-637.
	[ 马冬雪, 刘仁林 (2012). 天然群落枯枝落叶浸提液与其它处理对伯乐树种子发芽的比较研究. 林业科学研究, 25,632-637.]
[60]	Marques MCM, Burslem DFRP (2015). Multiple stage recruitment limitation and density dependence effects in two tropical forests. Plant Ecology, 216,1243-1255. DOI URL
[61]	Marteinsdóttir B (2014). Seed rain and seed bank reveal that seed limitation strongly influences plant community assembly in grasslands. PLOS ONE, 9,e103352. DOI: 10. 1371/journal.pone.0103352. DOI URL
[62]	Martinez-Ramos M, Alvarez-Buylla ER (1995). Seed dispersal and patch dynamics in tropical rain forests: a demographic approach. Écoscience, 2,223-229. DOI URL
[63]	Mayer R, Erschbamer B (2011). Seedling recruitment and seed-microsite limitation in traditionally grazed plant communities of the alpine zone. Basic and Applied Ecology, 12,10-20. DOI URL
[64]	McEuen AB, Curran LM (2004). Seed dispersal and recruitment limitation across spatial scales in temperate forest fragments. Ecology, 85,507-518. DOI URL
[65]	Mendoza I, Gómez-Aparicio L, Zamora R, Matías L (2009). Recruitment limitation of forest communities in a degraded Mediterranean landscape. Journal of Vegetation Science, 20,367-376. DOI URL
[66]	Molofsky J, Lanza J, Crone EE (2000). Plant litter feedback and population dynamics in an annual plant, Cardamine pensylvanica. Oecologia, 124,522-528.
[67]	Moreira X, Abdala-Roberts L, Zas R, Merlo E, Lombardero MJ, Sampedro L, Mooney KA (2016). Masting behaviour in a Mediterranean pine tree alters seed predator selection on reproductive output. Plant Biology, 18,973-980. DOI URL
[68]	Muller-Landau HC, Wright SJ, Calderón O, Hubbell SP, Foster RB (2002). Assessing recruitment limitation: concepts, methods and case-studies from a tropical forest//Levey DJ, Silva WR, Wallingford MG. Seed Dispersal and Frugivory: Ecology, Evolution and Conservation. CABI, Wallingford.35-53.
[69]	Münzbergová Z, Herben T (2005). Seed, dispersal, microsite, habitat and recruitment limitation: identification of terms and concepts in studies of limitations. Oecologia, 145,1-8. PMID
[70]	Overbeck G, Kiehl K, Abs C (2003). Seedling recruitment of Succisella inflexa in fen meadows: importance of seed and microsite availability. Applied Vegetation Science, 6,97-104. DOI URL
[71]	Palow DT, Oberbauer SF (2009). Soil type affects seedling shade response at low light for two Inga species from Costa Rica. Plant and Soil, 319,25-35. DOI URL
[72]	Pausas JG, Lamont BB, Paula S, Appezzato-Da-glória B, Fidelis A (2018). Unearthing belowground bud banks in fire-prone ecosystems. New Phytologist, 217,1435-1448. DOI URL
[73]	Putwain PD, Machin D, Harper JL (1968). Studies in the dynamics of plant populations: II. components and regulation of a natural population of Rumex acetosella L. Journal of Ecology, 56,421. DOI URL
[74]	Qian JQ, Wang ZW, Klimešová J, Lü X, Kuang WN, Liu ZM, Han XG (2017). Differences in below-ground bud bank density and composition along a climatic gradient in the temperate steppe of Northern China. Annals of Botany, 120,755-764. DOI URL
[75]	Rand TA, West NM, Russell FL, Louda SM (2020). Post-dispersal factors influence recruitment patterns but do not override the importance of seed limitation in populations of a native thistle. Oecologia, 193,143-153. DOI URL
[76]	Record S, Kobe RK, Vriesendorp CF, Finley AO (2016). Seedling survival responses to conspecific density, soil nutrients, and irradiance vary with age in a tropical forest. Ecology, 97,2406-2415. DOI PMID
[77]	Rey PJ, Ramírez JM, Sánchez-Lafuente AM (2006). Seed- vs. microsite-limited recruitment in a myrmecochorous herb. Plant Ecology, 184,213-222. DOI URL
[78]	Rosin C, Poulsen JR (2018). Seed traits, not density or distance from parent, determine seed predation and establishment in an Afrotropical forest. Biotropica, 50,881-888. DOI URL
[79]	Rother DC, Pizo MA, Siqueira T, Rodrigues RR, Jordano P (2015). Community-wide spatial and temporal discordances of seed-seedling shadows in a tropical rainforest. PLOS ONE, 10,e0123346. DOI: 10.1371/journal.pone.0123346. DOI URL
[80]	Russell ML, Vermeire LT (2015). Fire and nitrogen alter axillary bud number and activity in purple threeawn. Rangeland Ecology & Management, 68,65-70. DOI URL
[81]	Sagar GR, Harper JL (1960). Factors affecting the germination and early establishment of plantains (Plantago lanceolata, P. media and P. major)//Harper JL. The Biology of Weeds. Blackwell, Oxford.236-245.
[82]	Salazar A, Goldstein G, Franco AC, Miralles-Wilhelm F (2011). Timing of seed dispersal and dormancy, rather than persistent soil seed-banks, control seedling recruitment of woody plants in Neotropical savannas. Seed Science Research, 21,103-116. DOI URL
[83]	Schupp EW, Milleron T, Russo SE (2002). Dissemination limitation and the origin and maintenance of species-rich tropical forests//Levey DJ, Silva WR, Wallingford MG. Seed Dispersal and Frugivory: Ecology, Evolution and Conservation. CABI, Wallingford.19-33.
[84]	Seabloom EW (2011). Spatial and temporal variability in propagule limitation of California native grasses. Oikos, 120,291-301. DOI URL
[85]	Siemann E, Rogers WE (2006). Recruitment limitation, seedling performance and persistence of exotic tree monocultures. Biological Invasions, 8,979-991. DOI URL
[86]	Slodowicz D, Humbert JY, Arlettaz R (2019). The relative effectiveness of seed addition methods for restoring or re-creating species rich grasslands: a systematic review protocol. Environmental Evidence, 8, 28. DOI: 10.1186/s13750-019-0174-2. DOI
[87]	Solbreck C, Knape J (2017). Seed production and predation in a changing climate, new roles for resource and seed predator feedback? Ecology, 98,2301-2311. DOI URL
[88]	Titus JH, Bishop JG (2014). Propagule limitation and competition with nitrogen fixers limit conifer colonization during primary succession. Journal of Vegetation Science, 25,990-1003. DOI URL
[89]	Turnbull LA, Crawley MJ, Rees M (2000). Are plant populations seed-limited? A review of seed sowing experiments. Oikos, 88,225-238. DOI URL
[90]	Turnbull LA, Rees M, Crawley MJ (1999). Seed mass and the competition/colonization trade-off: a sowing experiment. Journal of Ecology, 87,899-912. DOI URL
[91]	Uriarte M, Bruna EM, Rubim P, Anciães M, Jonckheere I (2010). Effects of forest fragmentation on the seedling recruitment of a tropical herb: assessing seed vs. safe-site limitation. Ecology, 91,1317-1328. PMID
[92]	Urretavizcaya MF, Defossé GE (2013). Effects of nurse shrubs and tree shelters on the survival and growth of two Austrocedrus chilensis seedling types in a forest restoration trial in semiarid Patagonia, Argentina. Annals of Forest Science, 70,21-30. DOI URL
[93]	Vaughton G, Ramsey M (2001). Relationships between seed mass, seed nutrients, and seedling growth in Banksia cunninghamii (Proteaceae). International Journal of Plant Sciences, 162,599-606. DOI URL
[94]	Vilà M, Bartomeus I, Gimeno I, Traveset A, Moragues E (2006). Demography of the invasive geophyte Oxalis pes-caprae across a Mediterranean island. Annals of Botany, 97,1055-1062. DOI URL
[95]	Wang DL, Du J, Zhang BT, Ba L, Hodgkinson KC (2017). Grazing intensity and phenotypic plasticity in the clonal grass Leymus chinensis. Rangeland Ecology & Management, 70,740-747. DOI URL
[96]	Wang Z, Zhang GS, Wang LH, Hao YL, Wen GS (2005). Seed yield, seed bank and regeneration of natural Sabina vulgaris community in mu us sandland. Journal of Arid Land Resources and Environment, 19,195-200.
	[ 王哲, 张国盛, 王林和, 郝云龙, 温国胜 (2005). 毛乌素沙地天然臭柏群落种子产量、种子库及幼苗更新. 干旱区资源与环境, 19,195-200.]
[97]	Xiao ZS, Mi XC, Holyoak M, Xie WH, Cao K, Yang XF, Huang XQ, Krebs CJ (2017). Seed-predator satiation and Janzen-Connell effects vary with spatial scales for seed-feeding insects. Annals of Botany, 119,109-116. DOI URL
[98]	Xie TP, Zhang GF, Zhao ZG, Du GZ, He GY (2014). Intraspecific competition and light effect on reproduction of Ligularia virgaurea, an invasive native alpine grassland clonal herb. Ecology and Evolution, 4,817-825. DOI URL
[99]	Xu Y, Shen ZH, Li DX, Guo QF (2015). Pre-dispersal seed predation in a species-rich forest community: patterns and the interplay with determinants. PLOS ONE, 10,e0143040. DOI: 10.1371/journal.pone.0143040. DOI URL
[100]	Yang YF, Bai YP, Li JD (2012). Spatial difference and regularity of seed dispersal of Ulmus macrocarpa in Horqin Sandy Land, China. Chinese Journal of Plant Ecology, 36,747-753. DOI URL PMID
	[ 杨允菲, 白云鹏, 李建东 (2012). 科尔沁沙地黄榆种子散布的空间差异及规律. 植物生态学报, 36,747-753.] DOI PMID
[101]	Yang YF, Li JD (1996). Vegetative propagation of some rhizomatous grasses on Song-nen Plain and their population stability. Acta Prataculturae Sinica, 5,43-48.
	[ 杨允菲, 李建东 (1996). 松嫩平原几种根茎型禾草种群的营养繁殖特性及其持续更新分析. 草业学报, 5,43-48.]
[102]	Yang YF, Wei CY, Zhang BT, Liu B (2005). Dynamics of bud flow and bud bank of Phragmites communis population in dry land habitat of alkalinized meadow in the Songnen Plains of China. Chinese Journal of Applied Ecology, 16,854-858.
	[ 杨允菲, 魏春雁, 张宝田, 刘宝 (2005). 松嫩平原碱化草甸旱地生境芦苇种群的芽流和芽库动态. 应用生态学报, 16,854-858.]
[103]	Zárate DA, Andresen E, Santos-Heredia C (2019). Seed fate and seedling recruitment in monkey latrines in rustic cocoa plantations and rain forest in southern Mexico. Journal of Tropical Ecology, 35,18-25. DOI URL
[104]	Zhang HX, Zhou DW (2016). Current status in seed ecology. Pratacultural Science, 33,2221-2236.
	[ 张红香, 周道玮 (2016). 种子生态学研究现状. 草业科学, 33,2221-2236.]
[105]	Zhang JF, Li QH, Wang LL, Gao MY, Sun GJ, Xing L (2018). Effects of different nitrogen addition on bud bank traits and branching architecture of Nitraria tangutorum seedlings. Forest Research, 31,158-166.
	[ 张俊菲, 李清河, 王林龙, 高明远, 孙高洁, 邢磊 (2018). 白刺幼苗芽库及枝系构型对不同氮添加水平的响应. 林业科学研究, 31,158-166.]
[106]	Zhang YB, Li JW, Zhang H, Li JQ, Wu P, Zhao J (2005). Analysis on the factors cause the failure of Populus euhpratica sextual regeneration in nature. Science Technology and Engineering, 5,467-472.
	[ 张玉波, 李景文, 张昊, 李俊清, 乌平, 赵健 (2005). 额济纳胡杨有性繁殖失败因素分析. 科学技术与工程, 5,467-472.]
[107]	Zhao GJ, Xu XL, Ma HC, Huang D, Yang JJ, Shu YY, Ge L, Ping P (2016). Causes of difficulties with natural regeneration of a Bombax ceiba population in Hong-He dry-hot valleys (DHV). Acta Ecologica Sinica, 36,1342-1351.
	[ 赵高卷, 徐兴良, 马焕成, 黄冬, 杨建军, 舒媛媛, 葛娈, 平盼 (2016). 红河干热河谷木棉种群的天然更新. 生态学报, 36,1342-1351.]
[108]	Zhao LP, Tan ST, Bai X (2016). Effect of grazing and grazing exclusion management on offspring recruitment in a semiarid steppe on the Loess Plateau. Journal of Northwest A&F University(Natural Science Edition), 44(9),27-34.
	[ 赵凌平, 谭世图, 白欣 (2016). 封育和放牧对黄土高原典型草原繁殖更新的影响. 西北农林科技大学学报(自然科学版), 44(9),27-34.]
[109]	Zhou Y, Chen X, Wei XL, Wu Q, Li CC (2015). Effects of litter on the seedling regeneration and seed germination of Rhododendron agastum. Scientia Silvae Sinicae, 51(3),65-74.
	[ 周艳, 陈训, 韦小丽, 伍庆, 李朝婵 (2015). 凋落物对迷人杜鹃幼苗更新和种子萌发的影响. 林业科学, 51(3),65-74.]
[110]	Zimmermann J, Higgins SI, Grimm V, Hoffmann J, Münkemüller T, Linstädter A (2008). Recruitment filters in a perennial grassland: the interactive roles of fire, competitors, moisture and seed availability. Journal of Ecology, 96,1033-1044. DOI URL