深圳湾福田红树林自然保护区物种共存特征与物种分布概率

doi:10.17521/cjpe.2024.0323

摘要/Abstract

摘要：

深圳湾福田红树林自然保护区是我国唯一一处位于城市腹地的国家级自然保护区。受人为干扰较重的城市红树林湿地的潜在物种多样性的研究, 可为红树林生态湿地可持续管理提供路径。该研究对深圳湾福田红树林自然保护区不同区域(核心区/非核心区)进行生态学调查, 采用超几何法评估物种共存特征与物种分布概率, 并基于调查数据分析该区域的植被特征。主要结果有: (1)深圳湾福田红树林自然保护区现分布主要有9种红树植物, 秋茄树(Kandelia obovata)是优势种; 核心区中秋茄树重要值较大(平均81.2%), 在非核心区, 秋茄树重要值显著降低, 并出现了无瓣海桑(Sonneratia apetala)与海桑(S. caseolaris)。(2)海桑与无瓣海桑存在强的共存倾向, z分数值为2.82, 而无瓣海桑与秋茄树存在竞争排斥, z分数值为-2.41, z分数值反映了物种实际分布与随机预期的偏离程度。(3)现存物种多样性与群落完整性之间具有显著正相关关系, 群落完整性越高, 则现存物种多样性越丰富。非核心区部分样方的无瓣海桑分布概率较高, 面临扩散风险。核心区秋茄树的分布概率(0.51 ± 0.09)显著高于非核心区(0.41 ± 0.15), 但核心区部分样方(如8号样带的样方)群落完整性较低, 建议适当补种本土红树, 提高物种多样性。

关键词: 红树林, 物种共存, 物种库分布概率, 潜在物种多样性

Abstract:

Aims The Shenzhen Bay Mangrove Nature Reserve is the only national nature reserve in China located within an urban area. Studying the potential biodiversity of urban mangrove wetlands, which are significantly impacted by human disturbance, can provide a pathway for the sustainable management of mangrove ecosystems.

Methods This study conducted an ecological survey of different areas (core/non-core) in the Shenzhen Bay Mangrove National Nature Reserve. Species co-occurrence and distribution probability was assessed using the hypergeometric method, and the vegetation characteristics of the area were analyzed based on the survey data.

Important findings (1) A total of nine mangrove species were mainly distributed in the Shenzhen Bay Mangrove Nature Reserve, with Kandelia obovata being the dominant species. In the core area, K. obovata had the highest importance value (IV) (mean = 81.2%), while its importance value in the non-core area was significantly lower, where Sonneratia apetala and S. caseolaris appeared. (2) In the Shenzhen Bay mangrove ecosystem, S. caseolaris and S. apetala exhibited a strong coexistence tendency, with a z-score value of 2.82. Meanwhile, S. apetalaand K. obovata displayed competitive exclusion, with a z-score value of -2.41. z-score values reflect non-random species co-occurrence patterns. (3) A significant positive correlation was found between the observed species diversity and the community completeness index in the Shenzhen Bay mangroves. The higher the community completeness, the higher the observed species diversity. Some non-core area plots are at risk of S. apetala spreading, and ecosystem management and protection should be strengthened. The distribution probability of K. obovata in the core area (0.51 ± 0.09) is significantly higher than that in the non-core area (0.41 ± 0.15). However, in certain plots of the core area, such as the plots in the mid-tide zones of transects 8, community completeness is relatively low. It is recommended to appropriately replant native mangroves to enhance species diversity.

Key words: mangrove, species co-occurrence, distribution probability, potential biodiversity

郭欢敏, 沈小雪, 李瑞利. 深圳湾福田红树林自然保护区物种共存特征与物种分布概率. 植物生态学报, 2025, 49(11): 1833-1843. DOI: 10.17521/cjpe.2024.0323

GUO Huan-Min, SHEN Xiao-Xue, LI Rui-Li. Species co-occurrence and distribution probability in Futian Mangrove Nature Reserve of Shenzhen Bay. Chinese Journal of Plant Ecology, 2025, 49(11): 1833-1843. DOI: 10.17521/cjpe.2024.0323

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0323

https://www.plant-ecology.com/CN/Y2025/V49/I11/1833

图/表 9

图1 深圳湾福田红树林自然保护区采样点位置。1-10为样带编号。

Fig. 1 Location of sampling sites in Futian Mangrove Nature Reserve of Shenzhen Bay. 1-10 are belt transects ID.

图2 深圳湾福田红树林自然保护区非核心区与核心区的潜在物种多样性与现存物种多样性(平均值±标准误)。相同小写字母表示多样性指数在不同区域中无显著差异(p > 0.05)。

Fig. 2 Potential biodiversity and observed biodiversity in the non-core and core areas of Futian Mangrove Nature Reserve of Shenzhen Bay (mean ± SE). The same lowercase letters indicate no significant differences in diversity indices among different regions (p > 0.05).

图3 深圳湾福田红树林自然保护区群落完整性与物种多样性的相关性。

Fig. 3 Correlation between community completeness and species diversity in Futian Mangrove Nature Reserve of Shenzhen Bay.

图4 深圳湾福田红树林自然保护区非核心区与核心区的红树乔木的重要值(平均值±标准误)。不同小写字母表示同一物种在不同区域中分布概率值差异显著(p < 0.05)。Ac, 蜡烛果; Am, 海榄雌; Bg, 木榄; Bs, 海莲; Ko, 秋茄树; Sa, 无瓣海桑; Sc, 海桑。

Fig. 4 Important value of woody canopy mangrove plants in the non-core and core areas of Futian Mangrove Nature Reserve of Shenzhen Bay (mean ± SE). Different lowercase letters indicate significant differences in site-scale probability for the same species in the non-core and core areas (p < 0.05). Ac, Aegiceras corniculatum; Am, Avicennia marina; Bg, Bruguiera gymnorrhiza; Bs, B. sexangula; Ko, Kandelia obovata; Sa, Sonneratia apetala; Sc, S. caseolaris.

图5 深圳湾福田红树林自然保护区红树植物现存物种(A)、潜在物种(B)的分布概率值。每个格子代表一个样方, 灰色格子代表该物种在该样方中不存在(A)、存在(B)。Ac, 蜡烛果; Am, 海榄雌; Bg, 木榄; Bs, 海莲; Ko, 秋茄树; Sa, 无瓣海桑; Sc, 海桑。H、M、L分别代表高、中、低潮位样方。1-10为样带编号。

Fig. 5 Probability of observed (A), potential (B) species in site-specific species pool in Futian Mangrove Nature Reserve of Shenzhen Bay. Each grid represents one sampling site. Gray grid represents that the species is absent (A), observed (B) in the sampling site. Aa, Acrostichum aureum; Ac, Aegiceras corniculatum; Ai, Acanthus ilicifolius; Am, Avicennia marina; Bg, Bruguiera gymnorrhiza; Bs, B. sexangula; Ko, Kandelia obovata; Sa, Sonneratia apetala; Sc, S. caseolaris. H, M, and L denote sampling plots in the high, middle, and low tide zones, respectively. 1-10 are belt transects ID.

表1 深圳湾福田红树林自然保护区红树乔木的重要值(平均值±标准误)

Table 1 Importance values of mangrove plants in Futian Mangrove Nature Reserve of Shenzhen Bay (mean ± SE)

物种 Species	相对多度 Relative abundance (%)	相对频度 Relative frequency (%)	相对优势度 Relative dominance (%)	重要值 Importance value (%)
非核心区 Non-core area
无瓣海桑 Sonneratia apetala	33.1 ± 1.9	48.9 ± 6.6	18.3 ± 5.6	33.4 ± 4.7
海莲 Bruguiera sexangula	1.5 ± 0.9	0.3 ± 0.1	12.7 ± 2.2	4.8 ± 1.1
海桑 Sonneratia caseolaris	11.7 ± 3.6	19.9 ± 14.6	15.9 ± 5.7	15.8 ± 7.5
木榄 Bruguiera gymnorrhiza	1.7 ± 0.7	0.4 ± 0.2	19.6 ± 9.2	7.2 ± 3.3
蜡烛果 Aegiceras corniculatum	18.3 ± 20.7	2.6 ± 3.7	22.4 ± 8.9	14.4 ± 10.6
秋茄树 Kandelia obovata	57.9 ± 35.8	52.0 ± 30.9	30.2 ± 10.9	46.7 ± 25.1^b
海榄雌 Avicennia marina	8.0 ± 5.6	22.5 ± 10.4	20.5 ± 9.4	17.0 ± 7.0^a
核心区 Core area
木榄 Bruguiera gymnorrhiza	0.6 ± 0.0	0.9 ± 0.0	20.0 ± 0.0	7.2 ± 0.0
蜡烛果 Aegiceras corniculatum	9.8 ± 1.3	1.9 ± 0.5	41.7 ± 11.8	17.8 ± 4.5
秋茄树 Kandelia obovata	92.7 ± 5.3	97.6 ± 0.4	53.3 ± 5.8	81.2 ± 3.8^a
海榄雌 Avicennia marina	0.9 ± 0.3	1.2 ± 0.1	18.3 ± 2.4	6.8 ± 0.6^b

表2 深圳湾福田红树林自然保护区9种红树植物的分布概率值(平均值±标准误)

Table 2 Distribution probability of 9 mangrove species in Futian Mangrove Nature Reserve of Shenzhen Bay (mean ± SE)

物种 Species	分布概率 Site-scale probability
物种 Species	整个研究区 All sites	核心区 Core area	非核心区 Non-core area
无瓣海桑 Sonneratia apetala	0.30 ± 0.21	0.24 ± 0.12	0.33 ± 0.25
卤蕨 Acrostichum aureum	0.62 ± 0.11	0.69 ± 0.11^a	0.59 ± 0.10^b
海莲 Bruguiera sexangula	0.58 ± 0.12	0.62 ± 0.09	0.56 ± 0.13
海桑 Sonneratia caseolaris	0.37 ± 0.20	0.32 ± 0.14	0.40 ± 0.23
木榄 Bruguiera gymnorrhiza	0.65 ± 0.10	0.66 ± 0.07	0.64 ± 0.11
蜡烛果 Aegiceras corniculatum	0.57 ± 0.10	0.55 ± 0.09	0.58 ± 0.10
老鼠簕 Acanthus ilicifolius	0.50 ± 0.00	0.50 ± 0.00	0.50 ± 0.00
秋茄树 Kandelia obovata	0.44 ± 0.14	0.51 ± 0.09^a	0.41 ± 0.15^b
海榄雌 Avicennia marina	0.38 ± 0.07	0.37 ± 0.04	0.38 ± 0.07

图6 深圳湾福田红树林自然保护区非核心区与核心区的9种红树植物的分布概率(平均值±标准误)。不同小写字母表示同一物种在不同区域中分布概率差异显著(p < 0.05)。Aa, 卤蕨; Ac, 蜡烛果; Ai, 老鼠簕; Am, 海榄雌; Bg, 木榄; Bs, 海莲; Ko, 秋茄树; Sa, 无瓣海桑; Sc, 海桑。

Fig. 6 Distribution probability of 9 mangrove species in the non-core and core areas of Futian Mangrove Nature Reserve of Shenzhen Bay (mean ± SE). Different lowercase letters indicate significant differences in distribution probability for the same species in the non-core and core areas (p < 0.05). Aa, Acrostichum aureum; Ac, Aegiceras corniculatum; Ai, Acanthus ilicifolius; Am, Avicennia marina; Bg, Bruguiera gymnorrhiza; Bs, B. sexangula; Ko, Kandelia obovata; Sa, Sonneratia apetala; Sc, S. caseolaris.

图7 深圳湾福田红树林自然保护区9个红树物种两两之间的共存关系强弱。Aa, 卤蕨; Ac, 蜡烛果; Ai, 老鼠簕; Am, 海榄雌; Bg, 木榄; Bs, 海莲; Ko, 秋茄树; Sa, 无瓣海桑; Sc, 海桑。z分数值小于0表示负相关关系, 大于0为正相关关系, 等于0表示随机共存; 对角线的z分数无比较意义, 已用“-”标出。

Fig. 7 Relationship among of the 9 mangrove species in Futian Mangrove Nature Reserve of Shenzhen Bay. Aa, Acrostichum aureum; Ac, Aegiceras corniculatum; Ai, Acanthus ilicifolius; Am, Avicennia marina; Bg, Bruguiera gymnorrhiza; Bs, B. sexangula; Ko, Kandelia obovata; Sa, Sonneratia apetala; Sc, S. caseolaris. z-score values less than 0 indicate a negative correlation, values greater than 0 indicate a positive correlation, and values equal to 0 represent random cooccurrence. The z-scores along the diagonal are meaningless for comparison and have been marked with “-”.

参考文献 48

[1]	Analuddin K, Kadidae LO, Muhammad Yasir LO, Septiana A, Syahrir L, Rahim S, Pratama D, Fajar LA, Nadaoka K (2021). Blue carbon dynamics in mangroves and conservation of their services in the Coral Triangle Ecoregion, Southeast Sulawesi, Indonesia. Journal of Physics: Conference Series, 1899, 012016. DOI: 10.1088/1742-6596/1899/1/012016.
[2]	Arita HT (2016). Species co-occurrence analysis: pairwise versus matrix-level approaches. Global Ecology and Biogeography, 25, 1397-1400. DOI URL
[3]	Arrivabene HP, Campos CQ, da Costa Souza I, Wunderlin DA, Milanez CRD, Machado SR (2016). Differential bioaccumulation and translocation patterns in three mangrove plants experimentally exposed to iron. Consequences for environmental sensing. Environmental Pollution, 215, 302-313. DOI PMID
[4]	Branoff B (2002). Urban mangrove biology and ecology: emergent patterns and management implications// Makowski C, Finkl CW. Threats to Mangrove Forests: Hazards, Vulnerability, and Management. Springer. 521-537.
[5]	Carmona CP, Pärtel M (2021). Estimating probabilistic site-specific species pools and dark diversity from co-occurrence data. Global Ecology and Biogeography, 30, 316-326. DOI URL
[6]	Chen GG, Li YY, Cai LQ, Xu JJ, Lin PR, Chen WC, Huang SL (2017). Impacts of alien species Sonneratia apetala on morphological characteristics and biomass of native species Kandelia candel. Marine Sciences, 41(6), 26-33.
	[陈国贵, 李元跃, 蔡丽钦, 徐佳佳, 林沛然, 陈文城, 黄森磊 (2017). 红树植物外来种无瓣海桑对乡土种秋茄形态特征与生物量的影响. 海洋科学, 41(6), 26-33.]
[7]	Chen Q, Ma KM (2015). Research overview and trend on biological invasion in mangrove forests. Chinese Journal of Plant Ecology, 39, 283-299. DOI
	[陈权, 马克明 (2015). 红树林生物入侵研究概况与趋势. 植物生态学报, 39, 283-299.] DOI
[8]	Chen GZ, Mou SY, Zhang JH (1996). Ecologic study on the mangrove forest in Futian Nature Reserve, Shenzhen, China. Acta Scientiarum Naturalium Universitatis Sunyatseni, 35, 298-304.
	[陈桂珠, 缪绅裕, 章金鸿 (1996). 深圳福田红树林生态学研究. 中山大学学报(自然科学版), 35, 298-304.]
[9]	Chu CJ, Wang YS, Liu Y, Jiang L, He FL (2017). Advances in species coexistence theory. Biodiversity Science, 25, 345-354. DOI
	[储诚进, 王酉石, 刘宇, 蒋林, 何芳良 (2017). 物种共存理论研究进展. 生物多样性, 25, 345-354.] DOI
[10]	Dixon P (2003). VEGAN, a package of R functions for community ecology. Journal of Vegetation Science, 14, 927-930. DOI URL
[11]	Fang JY, Shen ZH, Tang ZY, Wang ZH (2004). The protocol for the survey plan for plant species diversity of China’s Mountains. Chinese Biodiversity, 12, 5-9.
	[方精云, 沈泽昊, 唐志尧, 王志恒 (2004). “中国山地植物物种多样性调查计划”及若干技术规范. 生物多样性, 12, 5-9.] DOI
[12]	Goldberg L, Lagomasino D, Thomas N, Fatoyinbo T (2020). Global declines in human-driven mangrove loss. Global Change Biology, 26, 5844-5855. DOI URL
[13]	Guan KL, Zhang XJ, Tan GW, Zeng F, Yi HL, Liao WB (2021). Species composition and structural diversity of typical mangrove communities in Shenzhen. Ecological Science, 40(4), 83-91.
	[关开朗, 张信坚, 谭广文, 曾凤, 易慧琳, 廖文波 (2021). 深圳市红树林典型群落的物种组成及结构多样性研究. 生态科学, 40(4), 83-91.]
[14]	Hagger V, Worthington TA, Lovelock CE, Adame MF, Amano T, Brown BM, Friess DA, Landis E, Mumby PJ, Morrison TH, O’Brien KR, Wilson KA, Zganjar C, Saunders MI (2022). Drivers of global mangrove loss and gain in social-ecological systems. Nature Communications, 13, 6373. DOI: 10.1038/s41467-022-33962-x. PMID
[15]	He ZY, Yen L, Huang HJ, Wang ZH, Zhao LL, Chen ZH, Lee SY, Peng YS (2022). Linkage between mangrove seedling colonization, sediment traits, and nitrogen input. Frontiers in Marine Science, 9, 793818. DOI: 10.3389/fmars.2022.793818.
[16]	Hu P, Xiang XL, Huang ZJ, Tan M, Yu SX, Yang Q (2024). Biomass dynamics in exotic and native mangrove species. Chinese Journal of Applied and Environmental Biology, 30, 929-934.
	[胡平, 向雪莲, 黄子健, 谭敏, 余世孝, 杨琼 (2024). 外来与乡土红树植物群落生物量动态. 应用与环境生物学报, 30, 929-934.]
[17]	Hu T, Chou QC, He SY, Xu HL, Shi XH (2016). Structural regulation and natural restoration of Futian mangrove in Shenzhen Bay. Chinese Journal of Ecology, 35, 1491-1496.
	[胡涛, 丑庆川, 何诗雨, 徐华林, 史秀华 (2016). 深圳湾红树林结构调控及自然恢复状况. 生态学杂志, 35, 1491-1496.]
[18]	Hu ZW, Wu JJ, Wang JZ, Zhang YH, Zhou HC, Gao CJ, Wang JJ, Wu GF (2023). How exotic Sonneratia species affect the spatiotemporal dynamics of mangroves in Shenzhen Bay, China: a remote sensing perspective. Ecological Indicators, 153, 110479. DOI: 10.1016/j.ecolind.2023.110479.
[19]	Jia MM, Wang ZM, Zhang YZ, Mao DH, Wang C (2018). Monitoring loss and recovery of mangrove forests during 42 years: the achievements of mangrove conservation in China. International Journal of Applied Earth Observation and Geoinformation, 73, 535-545. DOI URL
[20]	Lande R (1996). Statistics and partitioning of species diversity, and similarity among multiple communities. Oikos, 76, 5-13.
[21]	Lewis RJ, Szava-Kovats R, Pärtel M (2016). Estimating dark diversity and species pools: an empirical assessment of two methods. Methods in Ecology and Evolution, 7, 104-113. DOI URL
[22]	Li RL, Chai MW, Qiu GY, He B (2012). Mercury and copper accumulation during last fifty years and their potential ecological risk assessment in sediment of mangrove wetland of Shenzhen, China. Environmental Science, 33, 4276-4283.
	[李瑞利, 柴民伟, 邱国玉, 贺蓓 (2012). 近50年来深圳湾红树林湿地Hg、Cu累积及其生态危害评价. 环境科学, 33, 4276-4283.]
[23]	Li Z, Li Y, Zan QJ, Yu SX (2017). A comparison of mangrove community distribution and landscape pattern between Futian and Maipo Nature Reserve at Shenzhen Bay. Acta Scientiarum Naturalium Universitatis Sunyatseni, 56, 12-19.
	[李真, 李瑜, 昝启杰, 余世孝 (2017). 深圳福田与香港米埔红树林群落分布与景观格局比较. 中山大学学报(自然科学版), 56, 12-19.]
[24]	Liang SC, Liang MZ, Wu YL, Zan QJ, Wang YJ, Xie Q (2005). Analysis of the spatial structure of natural Sonneratia caseolaris + S. apetala forest in Futian, Shenzhen. Guihaia, 25, 393-398.
	[梁士楚, 梁铭忠, 吴苑玲, 昝启杰, 王勇军, 谢强 (2005). 深圳福田海桑+无瓣海桑自然林的空间结构分析. 广西植物, 25, 393-398.]
[25]	Liao BW, Li M, Zheng SF, Chen YJ, Zhong CR, Huang ZQ (2005). Niches of several mangrove species in Dongzhai Harbor of Hainan Island. Chinese Journal of Applied Ecology, 16, 403-407.
	[廖宝文, 李玫, 郑松发, 陈玉军, 钟才荣, 黄仲琪 (2005). 海南岛东寨港几种红树植物种间生态位研究. 应用生态学报, 16, 403-407.]
[26]	Liu BE, Liao BW (2013). The physio-ecological response of Acanthus ilicifolius seedlings to different degrees of light intensity in tide environment. Forest Research, 26, 192-199.
	[刘滨尔, 廖宝文 (2013). 老鼠簕幼苗在潮汐环境下对不同光强的生理生态响应. 林业科学研究, 26, 192-199.]
[27]	Ma ZS (2019). A new DTAR (diversity-time-area relationship) model demonstrated with the indoor microbiome. Journal of Biogeography, 46, 2024-2041. DOI URL
[28]	Moeslund JE, Brunbjerg AK, Clausen KK, Dalby L, Fløjgaard C, Juel A, Lenoir J (2017). Using dark diversity and plant characteristics to guide conservation and restoration. Journal of Applied Ecology, 54, 1730-1741. DOI URL
[29]	Pärtel M, Szava-Kovats R, Zobel M (2011). Dark diversity: shedding light on absent species. Trends in Ecology & Evolution, 26, 124-128. DOI URL
[30]	Pärtel M, Szava-Kovats R, Zobel M (2013). Community completeness: linking local and dark diversity within the species pool concept. Folia Geobotanica, 48, 307-317. DOI URL
[31]	Peng D, Chen LZ, Pennings SC, Zhang YH (2018). Using a marsh organ to predict future plant communities in a Chinese estuary invaded by an exotic grass and mangrove. Limnology and Oceanography, 63, 2595-2605. DOI URL
[32]	Peng X, Jin GZ (2022). Effects of plant characteristics and environmental factors on the dark diversity in a broadleaved Korean pine forest. Chinese Journal of Plant Ecology, 46, 656-666. DOI
	[彭鑫, 金光泽 (2022). 植物特性和环境因子对阔叶红松林暗多样性的影响. 植物生态学报, 46, 656-666.] DOI
[33]	Ren H, Jian SG, Lu HF, Zhang QM, Shen WJ, Han WD, Yin ZY, Guo QF (2008). Restoration of mangrove plantations and colonization by native species in Leizhou bay, South China. Ecological Research, 23, 401-407. DOI URL
[34]	Ren H, Lu HF, Shen WJ, Huang C, Guo QF, Li ZA, Jian SG (2009). Sonneratia apetala Buch.Ham in the mangrove ecosystems of China: an invasive species or restoration species? Ecological Engineering, 35, 1243-1248. DOI URL
[35]	Ren H, Wu XM, Ning TZ, Huang G, Wang J, Jian SG, Lu HF (2011). Wetland changes and mangrove restoration planning in Shenzhen Bay, Southern China. Landscape and Ecological Engineering, 7, 241-250. DOI URL
[36]	Simpson EH (1949). Measurement of diversity. Nature, 163, 688. DOI: 10.1038/163688a0.
[37]	Smart LS, Vukomanovic J, Sills EO, Sanchez G (2021). Cultural ecosystem services caught in a ‘coastal squeeze’ between sea level rise and urban expansion. Global Environmental Change, 66, 102209. DOI: 10.1016/j.gloenvcha.2020.102209.
[38]	Song SS, Ding YL, Li W, Meng YC, Zhou J, Gou RK, Zhang CH, Ye SB, Saintilan N, Krauss KW, Crooks S, Lv SG, Lin GH (2023). Mangrove reforestation provides greater blue carbon benefit than afforestation for mitigating global climate change. Nature Communications, 14, 756. DOI: 10.1038/s41467-023-36477-1.
[39]	Tang LL, Wang RX, He KS, Shi C, Yang T, Huang YP, Zheng PF, Shi FC (2019). Throwing light on dark diversity of vascular plants in China: predicting the distribution of dark and threatened species under global climate change. PeerJ, 7, e6731. DOI: 10.7717/peerj.6731.
[40]	Toral-Granda MV, Causton CE, Jäger H, Trueman M, Izurieta JC, Araujo E, Cruz M, Zander KK, Izurieta A, Garnett ST (2017). Alien species pathways to the Galapagos Islands, Ecuador. PLoS ONE, 12, e0184379. DOI: 10.1371/journal.pone.0184379.
[41]	Trindade DPF, Carmona CP, Pärtel M (2020). Temporal lags in observed and dark diversity in the Anthropocene. Global Change Biology, 26, 3193-3201. DOI PMID
[42]	Ulrich W, Gotelli NJ (2013). Pattern detection in null model analysis. Oikos, 122, 2-18. DOI URL
[43]	Veech JA (2013). A probabilistic model for analysing species co-occurrence. Global Ecology and Biogeography, 22, 252-260. DOI URL
[44]	Wang WQ, Shi JB, Chen LZ (2021). Research of Conservation and Restoration Strategy of Mangrove Wetlands in China. China Environmental Science Press, Beijing.
	[王文卿, 石建斌, 陈鹭真 (2021). 中国红树林湿地保护与恢复战略研究. 中国环境出版集团, 北京.]
[45]	Wu QH, Leung JYS, Yuan X, Huang XX, Li HY, Huang ZY, Li Y (2015). Biological risk, source and pollution history of organochlorine pesticides (OCPs) in the sediment in Nansha mangrove, South China. Marine Pollution Bulletin, 96, 57-64. DOI PMID
[46]	Zan QJ, Wang BS, Wang YJ, Li MG (2003). Ecological assessment on the introduced Sonneratia caseolaris and S. apetala at the mangrove forest of Shenzhen Bay, China. Acta Botanica Sinica, 544-551.
	[昝启杰, 王伯荪, 王勇军, 李鸣光 (2003). 深圳湾红树林引种海桑、无瓣海桑的生态评价. 生态学报, 544-551.]
[47]	Zhang LE, Liao BW, Guan W (2011). Effects of simulated tide inundation on seed germination and seedling growth of mangrove species Acanthus ilicifolius. Chinese Journal of Ecology, 30, 2165-2172.
	[张留恩, 廖宝文, 管伟 (2011). 模拟潮汐淹浸对红树植物老鼠簕种子萌发及幼苗生长的影响. 生态学杂志, 30, 2165-2172.]
[48]	Zhang YQ, Zhang Z, Jiang BQ, Shen XX, Li RL (2022). Ecosystem health assessment and management strategies of urban mangrove: a case study of Guangdong-Hong Kong-Macao Greater Bay Area. China Environmental Science, 42, 2352-2369.
	[张月琪, 张志, 江鎞倩, 沈小雪, 李瑞利 (2022). 城市红树林生态系统健康评价与管理对策——以粤港澳大湾区为例. 中国环境科学, 42, 2352-2369.]