Differential responses of wetland plant species to water depth changes and litter cover

doi:10.17521/cjpe.2025.0014

Abstract

Abstract:

Aims Water depth is a critical environmental factor in wetland ecosystems, influencing plant growth, regeneration, and distribution patterns. As an essential component of natural ecosystems, litter affects the regeneration of plant seedlings through various mechanisms such as light attenuation, mechanical obstruction, and allelopathy. Wetland ecosystems tend to accumulate a larger amount of litter because of flooding and anaerobic conditions. However, few studies have investigated the interactive effects of litter cover and water depth on the regeneration of plant seedlings.

Methods This research simulated the field environmental conditions in mobile rain shelters to analyze the influences of water depth (wet, water-saturated, and flooded) and litter cover (0, 500, 1 000, and 1 500 g·m^-2) on the seed germination rates and emergence rates of Echinochloa crus-galli, E. crus-gallivar. mitis, Bidens pilosa, and Alisma plantago-aquatica.

Important findings We found that the seed germination rates and emergence rates of E. crus-galli, E. crus-gallivar. mitis and A. plantago-aquatica were significantly affected by litter cover, water depth, and their interactions, while B. pilosa was only significantly influenced by litter cover. Litter cover decreased the germination and emergence rates of B. pilosa by 59.4% to 96.9% and 94.4% to 100.0%, respectively. The germination rate of B. pilosa seeds beneath litter was significantly reduced by 94% compared to that above litter, indicating that the shading effect resulting from litter cover was the primary cause restricting the germination of B. pilosa seeds. The germination rate of A. plantago-aquatica seeds was nearly zero under wet conditions, but increased to over 30% in water-saturated or flooded environments, suggesting that sufficient moisture is of vital importance for the germination of A. plantago-aquatica seeds. Under water-saturated or flooded conditions, 29% to 60% of the seeds of A. plantago-aquatica germinated when covered by litter. However, only 1.5% to 17% of the seedlings penetrated through the litter layer. We posit that mechanical obstruction caused by litter cover is a critical factor influencing the seedling regeneration of A. plantago-aquatica. This study can help understand the distribution patterns and succession dynamics of wetland vegetation under environmental changes and has important implications for the ecological restoration and scientific management of wetland vegetation.

Key words: litter, water depth, wetland ecosystem, seedling germination

ZHANG Xu-Dong, LIU Bo, ZHANG Dan, WU Hai-Tao, PAN Yuan, ZHENG Hao-Wen, LI Rui, YAN Shuo, SHEN Min-Yan, LAI Ming-Zi. Differential responses of wetland plant species to water depth changes and litter cover[J]. Chin J Plant Ecol, 2025, 49(12): 2069-2079.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2025.0014

https://www.plant-ecology.com/EN/Y2025/V49/I12/2069

Figures/Tables 5

References 56

[1]	Berg B, Staaf H (1981). Leaching, accumulation and release of nitrogen in decomposing forest litter. Ecological Bulletins, 163-178.
[2]	Bonanomi G, Incerti G, Abd El-Gawad AM, Cesarano G, Sarker TC, Saulino L, Lanzotti V, Saracino A, Rego FC, Mazzoleni S (2018). Comparing chemistry and bioactivity of burned vs. decomposed plant litter: different pathways but same result? Ecology, 99, 158-171. DOI PMID
[3]	Bosy JL, Reader RJ (1995). Mechanisms underlying the suppression of forb seedling emergence by grass (Poa pratensis) litter. Functional Ecology, 9, 635-639. DOI URL
[4]	Cai RR, Liu YG, Shen LD, Zheng XB, Dai GH (2024). A dataset of litter recovery amount and stock dynamics in broad-leaved Korean pine forests in Changbai Mountain (2011-2020). China Scientific Data, 9(4), 222-230.
	[蔡榕榕, 刘雅各, 沈历都, 郑兴波, 戴冠华 (2024). 2011-2020年长白山阔叶红松林凋落物回收量和现存量动态数据集. 中国科学数据, 9(4), 222-230.]
[5]	Campbell G, Lambert JDH, Arnason T, Neil Towers GH (1982). Allelopathic properties of α-terthienyl and phenylheptatriyne, naturally occurring compounds from species of Asteraceae. Journal of Chemical Ecology, 8, 961-972. DOI PMID
[6]	Carson WP, Peterson CJ (1990). The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance. Oecologia, 85, 8-13. DOI PMID
[7]	Castle S, Rejmánková E, Foley J, Parmenter S (2019). Hydrologic alterations impact plant litter decay rate and ecosystem resilience in Mojave wetlands. Restoration Ecology, 27, 1094-1104. DOI URL
[8]	Cerdà A, Doerr SH (2008). The effect of ash and needle cover on surface runoff and erosion in the immediate post-fire period. Catena, 74, 256-263. DOI URL
[9]	Deba F, Xuan TD, Yasuda M, Tawata S (2007). Herbicidal and fungicidal activities and identification of potential phytotoxins from Bidens pilosa L. var. radiata Scherff. Weed Biology and Management, 7, 77-83. DOI URL
[10]	Ding JP, Luo YQ (2022). Effect of litter water extraction of Artemisia halodendron on seed germination of four forage species. Environmental Ecology, 4(8), 40-46.
	[丁杰萍, 罗永清 (2022). 差巴嘎蒿凋落物水浸提液对4种牧草种子萌发的影响研究. 环境生态学, 4(8), 40-46.]
[11]	Facelli JM, Pickett STA (1991). Plant litter: Light interception and effects on an old-field plant community. Ecology, 72, 1024-1031. DOI URL
[12]	Facelli JM, Williams R, Fricker S, Ladd B (1999). Establishment and growth of seedlings of Eucalyptus obliqua: interactive effects of litter, water, and pathogens. Australian Journal of Ecology, 24, 484-494. DOI URL
[13]	Fahey C, Angelini C, Flory SL (2018). Grass invasion and drought interact to alter the diversity and structure of native plant communities. Ecology, 99, 2692-2702. DOI PMID
[14]	Fay PA, Guntenspergen GR, Olker JH, Johnson WC (2016). Climate change impacts on freshwater wetland hydrology and vegetation cover cycling along a regional aridity gradient. Ecosphere, 7, e01504. DOI: 10.1002/ecs2.1504.
[15]	Geißler C, Kühn P, Böhnke M, Bruelheide H, Shi X, Scholten T (2012). Splash erosion potential under tree canopies in subtropical SE China. Catena, 91, 85-93. DOI URL
[16]	Gong YW, Wang L, Tan Y, Chen G, Zhang ZX, Yang HJ (2021). Distribution characteristics and seasonal dynamics of litter in a headwater stream bed of the Changbai Mountain. Ecological Science, 40(2), 74-81.
	[宫雨薇, 王璐, 谭颖, 陈鸽, 张振兴, 杨海军 (2021). 长白山源头溪流河床凋落物的分布特征及季节动态. 生态科学, 40(2), 74-81.]
[17]	Grice AC, Vanderduys EP, Perry JJ, Cook GD (2013). Patterns and processes of invasive grass impacts on wildlife in Australia. Wildlife Society Bulletin, 37, 478-485.
[18]	Hodkinson DJ, Askew AP, Thompson K, Hodgson JG, Bakker JP, Bekker RM (1998). Ecological correlates of seed size in the British flora. Functional Ecology, 12, 762-766. DOI URL
[19]	Ibáñez I, Schupp EW (2002). Effects of litter, soil surface conditions, and microhabitat on Cercocarpus ledifolius Nutt.: seedling emergence and establishment. Journal of Arid Environments, 52, 209-221. DOI URL
[20]	Jiang YX (1981). Phytocenological role of forest floor in subalpine fir forests in western Sichuan Province. Acta Phytoecologia et Geobotanica Sinica, 5(2), 89-98.
	[蒋有绪 (1981). 川西亚高山冷杉林枯枝落叶层的群落学作用. 植物生态学与地植物学丛刊, 5(2), 89-98.]
[21]	Junk WJ, Piedade MTF, Schöngart J, Cohn-Haft M, Adeney JM, Wittmann F (2011). A classification of major naturally-occurring Amazonian lowland wetlands. Wetlands, 31, 623-640. DOI URL
[22]	Kettenring KM, Tarsa EE (2020). Need to seed? Ecological, genetic, and evolutionary keys to seed-based wetland restoration. Frontiers in Environmental Science, 8, 109. DOI: 10.3389/fenvs.2020.00109.
[23]	Kuiters AT (1990). Role of phenolic substances from decomposing forest litter in plant-soil interactions. Acta Botanica Neerlandica, 39, 329-348. DOI URL
[24]	Li AG, Lin CF, Hu MY, Liu XF, Song HW, Zhang L, Yang YS (2022). Effects of warming on physicochemical property of Cunninghamia lanceolata branch and leaf litter in subtropical plantation. Chinese Journal of Applied Ecology, 33, 2711-2717.
	[李澳归, 林成芳, 胡明艳, 刘小飞, 宋豪威, 张磊, 杨玉盛 (2022). 增温对亚热带杉木枝和叶凋落物理化性质的影响. 应用生态学报, 33, 2711-2717.] DOI
[25]	Li SF, Xu QY, Zhang JE, Ye YQ (2020). Effect of continuous overlay mulching of rice straw-plastic film on control of invasive plant Bidens pilosa L. Chinese Journal of Eco-Agriculture, 28, 1682-1691.
	[李赛飞, 许秋园, 章家恩, 叶延琼 (2020). 水稻秸秆-薄膜连续叠加覆盖对三叶鬼针草的控制效果. 中国生态农业学报(中英文), 28, 1682-1691.]
[26]	Li W, Tan R, Yang YM, Wang J (2014). Plant diversity as a good indicator of vegetation stability in a typical plateau wetland. Journal of Mountain Science, 11, 464-474. DOI URL
[27]	Li XF, Xu X, Wang BX, Huang YY, Wang ZF, Li JY (2012). Effects of forest litter layer on regeneration of Populus cathayana natural population in Xiaowutai Mountains in China. Chinese Journal of Plant Ecology, 36, 109-116. DOI
	[李霄峰, 胥晓, 王碧霞, 黄尤优, 王志峰, 李俊钰 (2012). 小五台山森林落叶层对天然青杨种群更新方式的影响. 植物生态学报, 36, 109-116.] DOI
[28]	Liu JY, Ding CJ, Zhang WX, Wei YW, Zhou YB, Zhu WX (2022). Litter mixing promoted decomposition rate through increasing diversities of phyllosphere microbial communities. Frontiers in Microbiology, 13, 1009091. DOI: 10.3389/fmicb.2022.1009091.
[29]	Liu Q, Liu JL, Liu HF, Liang LQ, Cai YP, Wang X, Li CH (2020). Vegetation dynamics under water-level fluctuations: implications for wetland restoration. Journal of Hydrology, 581, 124418. DOI: 10.1016/j.cropro.2021.105743.
[30]	Monroy S, Larrañaga A, Martínez A, Pérez J, Molinero J, Basaguren A, Pozo J (2023). Temperature sensitivity of microbial litter decomposition in freshwaters: role of leaf litter quality and environmental characteristics. Microbial Ecology, 85, 839-852. DOI
[31]	Moravcová L, Zákravský P, Hroudová Z (2001). Germination and seedling establishment in Alisma gramineum, A. plantago-aquatica and A. lanceolatum under different environmental conditions. Folia Geobotanica, 36, 131-146. DOI URL
[32]	Mulhouse JM, de Steven D, Lide RF, Sharitz RR (2005). Effects of dominant species on vegetation change in Carolina bay wetlands following a multi-year drought. Journal of the Torrey Botanical Society, 132, 411-420. DOI URL
[33]	Petraglia A, Cacciatori C, Chelli S, Fenu G, Calderisi G, Gargano D, Abeli T, Orsenigo S, Carbognani M (2019). Litter decomposition: effects of temperature driven by soil moisture and vegetation type. Plant and Soil, 435, 187-200. DOI
[34]	Raulings EJ, Morris K, Roache MC, Boon PI (2010). The importance of water regimes operating at small spatial scales for the diversity and structure of wetland vegetation. Freshwater Biology, 55, 701-715. DOI URL
[35]	Ren YL, Lu M, Fan FX, Peng SX (2019). The relationship between soil fungi and physico-chemical properties in swamp meadow of plateau wetlands. Ecological Science, 38(1), 42-49.
	[任玉连, 陆梅, 范方喜, 彭淑娴 (2019). 高原湿地沼泽化草甸土壤真菌与理化性质的关系. 生态科学, 38(1), 42-49.]
[36]	Rezvani M, Nadimi S, Zaefarian F, Chauhan BS (2021). Environmental factors affecting seed germination and seedling emergence of three Phalaris species. Crop Protection, 148, 105743. DOI: 10.1016/j.cropro.2021.105743.
[37]	Rosbakh S, Phartyal SS, Poschlod P (2020). Seed germination traits shape community assembly along a hydroperiod gradient. Annals of Botany, 125, 67-78. DOI PMID
[38]	Ruprecht E, Szabó A (2012). Grass litter is a natural seed trap in long-term undisturbed grassland. Journal of Vegetation Science, 23, 495-504. DOI URL
[39]	Seabloom EW, van der Valk AG, Moloney KA (1998). The role of water depth and soil temperature in determining initial composition of prairie wetland coenoclines. Plant Ecology, 138, 203-216. DOI
[40]	Song HL, Sun ZG, Sun JK, Mou XJ, Jiang HH, Sun WG (2012). Effects of nitrogen and phosphorus on seed germination and seedling growth of Suaeda salsa under different growth conditions of the Yellow River Estuary. Acta Prataculturae Sinica, 21(6), 30-41.
	[宋红丽, 孙志高, 孙景宽, 牟晓杰, 姜欢欢, 孙文广 (2012). 氮、磷输入对黄河口潮滩湿地不同生境下碱蓬种子萌发与幼苗生长的影响. 草业学报, 21(6), 30-41.]
[41]	Unsworth RKF, Rees SC, Bertelli CM, Esteban NE, Furness EJ, Walter B (2022). Nutrient additions to seagrass seed planting improve seedling emergence and growth. Frontiers in Plant Science, 13, 1013222. DOI: 10.3389/fpls.2022.1013222.
[42]	Vazquez-Yanes C, Orozco-Segovia A, Rincon E, Sanchez-Coronado ME, Huante P, Toledo JR, Barradas VL (1990). Light beneath the litter in a tropical forest: effect on seed germination. Ecology, 71, 1952-1958. DOI URL
[43]	Wang P, Li XY, Re FK, Lin LS, Zeng FJ (2024). A dataset of monthly litter recovery from the desert grassland in Southern Tarim Basin during 2010-2020. China Scientific Data, 9(2), 232-242.
	[王鹏, 李向义, 热甫开提, 林丽莎, 曾凡江 (2024). 2010-2020年塔里木盆地南部荒漠草地凋落物回收量月动态数据集. 中国科学数据, 9(2), 232-242.]
[44]	Wu HT, Jiang M, Lü XG, Mao DH, Shen XJ, Wang GD (2022). Basic theories and key technical problems of wetlands protection and restoration——The 289th Shuangqing Forum. Wetland Science, 20, 128-132.
	[武海涛, 姜明, 吕宪国, 毛德华, 神祥金, 王国栋 (2022). 湿地保护和修复的基础理论及关键技术问题——第289期双清论坛. 湿地科学, 20, 128-132.]
[45]	Xiao C, Wang XF, Xia J, Liu GH (2010). The effect of temperature, water level and burial depth on seed germination of Myriophyllum spicatum and Potamogeton malaianus. Aquatic Botany, 92, 28-32. DOI URL
[46]	Xie XG, Chen Y, Bu YQ, Dai CC (2014). A review of allelopathic researches on phenolic acids. Acta Ecologica Sinica, 34, 6417-6428.
	[谢星光, 陈晏, 卜元卿, 戴传超 (2014). 酚酸类物质的化感作用研究进展. 生态学报, 34, 6417-6428.]
[47]	Xiong SJ, Johansson ME, Hughes FMR, Hayes A, Richards KS, Nilsson C (2003). Interactive effects of soil moisture, vegetation canopy, plant litter and seed addition on plant diversity in a wetland community. Journal of Ecology, 91, 976-986. DOI URL
[48]	Yan PF, Zhan PF, Xiao DR, Wang Y, Yu R, Liu ZY, Wang H (2019). Effects of simulated warming and decomposition interface on the litter decomposition rate of Zizania latifolia and its phyllospheric microbial community structure and function. Chinese Journal of Plant Ecology, 43, 107-118. DOI URL
	[闫鹏飞, 展鹏飞, 肖德荣, 王燚, 余瑞, 刘振亚, 王行 (2019). 模拟增温及分解界面对茭草凋落物分解速率及叶际微生物结构和功能的影响. 植物生态学报, 43, 107-118.] DOI
[49]	Yasin M, Andreasen C (2016). Effect of reduced oxygen concentration on the germination behavior of vegetable seeds. Horticulture Environment, and Biotechnology, 57, 453-461. DOI URL
[50]	Yu J, Sharpe SM, Boyd NS (2020). Germination and emergence of common beggar’s-tick (Bidens alba) seeds at two different stages of afterripening as affected by environmental factors. Weed Science, 68, 503-509. DOI URL
[51]	Yue K, Ni X, Fornara DA, Peng Y, Liao S, Tan S, Wang D, Wu F, Yang Y (2021). Dynamics of calcium, magnesium, and manganese during litter decomposition in alpine forest aquatic and terrestrial ecosystems. Ecosystems, 24, 516-529. DOI
[52]	Zhang JJ, Li XZ, Wang YN, Deng JJ, Zhou L, Zhou WM, Yu DP, Wang QW (2024). Research advance in effects of solar radiation on litter decomposition in terrestrial ecosystems. Chinese Journal of Applied Ecology, 35, 2463-2472.
	[张娟娟, 李星志, 王亚楠, 邓娇娇, 周莉, 周旺明, 于大炮, 王庆伟 (2024). 太阳辐射对陆地生态系统凋落物分解影响的研究进展. 应用生态学报, 35, 2463-2472.] DOI
[53]	Zhang X, Ni X, Heděnec P, Yue K, Wei X, Yang J, Wu FZ (2022). Litter facilitates plant development but restricts seedling establishment during vegetation regeneration. Functional Ecology, 36, 3134-3147. DOI URL
[54]	Zhang Z, Wang H, Cao J, Li G, Chauhan BS (2023). Seed biology of alkali barnyardgrass (Echinochloa crus-galli var. zelayensis) and junglerice (Echinochloa colona) for improved management in direct-seeded rice. Weed Science, 71, 112-123. DOI URL
[55]	Zhao C, Cai YB, Huang X, Liu QQ, Zhu CX, Yu JD, Wang ZN, Liu B (2019). Effects of Chinese fir litter cover on its seedling emergence and early growth. Chinese Journal of Applied Ecology, 30, 481-488. DOI
	[赵冲, 蔡一冰, 黄晓, 刘青青, 朱晨曦, 于姣妲, 王正宁, 刘博 (2019). 杉木凋落物覆盖对自身幼苗出土和早期生长的影响. 应用生态学报, 30, 481-488.]
[56]	Zhu J, Sun G, Li W, Zhang Y, Miao G, Noormets A, Mcnulty SG, King JS, Kumar M, Wang X (2017). Modeling the potential impacts of climate change on the water table level of selected forested wetlands in the southeastern United States. Hydrology and Earth System Sciences, 21, 6289-6305. DOI URL

物种 Species	因素 Factor	自由度 df	F	p
稗 Echinochloa crus-galli	凋落物量 Litter amount	3	44.342	<0.001
	水深 Water depth	2	101.853	<0.001
	凋落物量×水深 Litter amount × water depth	6	12.195	<0.001
无芒稗 Echinochloa crus-galli var. mitis	凋落物量 Litter amount	3	34.152	<0.001
	水深变化 Water depth	2	19.946	<0.001
	凋落物量×水深 Litter amount × water depth	6	4.268	0.002
鬼针草 Bidens pilosa	凋落物量 Litter amount	3	287.734	<0.001
	水深变化 Water depth	2	2.549	0.092
	凋落物量×水深 Litter amount × water depth	6	2.266	0.059
泽泻 Alisma plantago-aquatica	凋落物量 Litter amount	3	9.995	<0.001
	水深变化 Water depth	2	198.117	<0.001
	凋落物量×水深 Litter amount × water depth	6	3.312	0.011

物种 Species	因素 Factor	自由度 df	F	p
稗 Echinochloa crus-galli	凋落物量 Litter amount	3	44.342	<0.001
	水深 Water depth	2	101.853	<0.001
	凋落物量×水深 Litter amount × water depth	6	12.195	<0.001
无芒稗 Echinochloa crus-galli var. mitis	凋落物量 Litter amount	3	34.152	<0.001
	水深变化 Water depth	2	19.946	<0.001
	凋落物量×水深 Litter amount × water depth	6	4.268	0.002
鬼针草 Bidens pilosa	凋落物量 Litter amount	3	287.734	<0.001
	水深变化 Water depth	2	2.549	0.092
	凋落物量×水深 Litter amount × water depth	6	2.266	0.059
泽泻 Alisma plantago-aquatica	凋落物量 Litter amount	3	9.995	<0.001
	水深变化 Water depth	2	198.117	<0.001
	凋落物量×水深 Litter amount × water depth	6	3.312	0.011

物种 Species	因素 Factor	自由度 df	F	p
稗 Echinochloa crus-galli	凋落物量 Litter amount	3	143.965	<0.001
	水深 Water depth	2	84.980	<0.001
	凋落物量×水深 Litter amount × water depth	6	11.317	<0.001
无芒稗 Echinochloa crus-galli var. mitis	凋落物量 Litter amount	3	159.128	<0.001
	水深 Water depth	2	8.464	<0.001
	凋落物量×水深 Litter amount × water depth	6	2.654	0.031
鬼针草 Bidens pilosa	凋落物量 Litter amount	3	652.393	<0.001
	水深 Water depth	2	2.447	0.101
	凋落物量×水深 Litter amount × water depth	6	2.169	0.069
泽泻 Alisma plantago-aquatica	凋落物量 Litter amount	3	68.739	<0.001
	水深 Water depth	2	35.085	<0.001
	凋落物量×水深 Litter amount × water depth	6	13.205	<0.001

物种 Species	因素 Factor	自由度 df	F	p
稗 Echinochloa crus-galli	凋落物量 Litter amount	3	143.965	<0.001
	水深 Water depth	2	84.980	<0.001
	凋落物量×水深 Litter amount × water depth	6	11.317	<0.001
无芒稗 Echinochloa crus-galli var. mitis	凋落物量 Litter amount	3	159.128	<0.001
	水深 Water depth	2	8.464	<0.001
	凋落物量×水深 Litter amount × water depth	6	2.654	0.031
鬼针草 Bidens pilosa	凋落物量 Litter amount	3	652.393	<0.001
	水深 Water depth	2	2.447	0.101
	凋落物量×水深 Litter amount × water depth	6	2.169	0.069
泽泻 Alisma plantago-aquatica	凋落物量 Litter amount	3	68.739	<0.001
	水深 Water depth	2	35.085	<0.001
	凋落物量×水深 Litter amount × water depth	6	13.205	<0.001