钢—混立方体人工藻礁附着生物的定殖效果及其群落演替特征研究

李璐瑶; 李文涛; 李艳平; 赵祺; 张沛东

钢—混立方体人工藻礁附着生物的定殖效果及其群落演替特征研究

中国海洋大学海水养殖教育部重点实验室，青岛 266003

基金项目: 国家重点研发计划课题（2023YFD2401101）。

详细信息

作者简介:
李璐瑶（2000—），女，山东临沂人，主要从事海藻场退化机制与修复理论。 E-mail：Liluyao20321@163.com

通讯作者:
张沛东（1975—），男，教授；研究方向为海洋牧场生境构建理论与技术。E-mail：zhangpdsg@ouc.edu.cn

中图分类号: S953.1
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出版历程
- 收稿日期: 2026-02-04
- 修回日期: 2026-04-27
- 网络出版日期: 2026-05-14

Study on the colonization dynamics and successional trajectories of sessile organisms on steel-concrete cubic artificial algal reefs

The Key Laboratory of Mariculture (Ocean University of China), Ministry of Education, Qingdao, Shandong 266003, China

摘要

摘要: 针对海藻场修复中人工藻礁单一材料生境异质性不足、生物附着效果较差的问题，本研究设计了一种钢材—混凝土异质型复合材料藻礁，制作礁体12个并于2021年8月投放至荣成马山里海域，于2021年11月、2022年6月和9月及2023年5月和8月，分别对钢材和混凝土大型附着藻类和附着动物以及投礁区关键环境因子进行了跟踪监测。结果显示：（1）两种材料对大型藻类的建植效果存在显著不同，其中混凝土的大型藻类生物量显著高于钢材，整个监测周期平均达到钢材的1.7倍（P<0.05）；（2）大型附着藻类群落结构主要受季节和礁龄的双重驱动，呈现出由绿藻门向褐藻门的演替规律，红藻门主要出现在演替的过渡期但未形成稳定的优势阶段，在投礁后20个月形成了以裙带菜（Undaria pinnatifida）为绝对优势种（Y=0.99）的稳定顶级群落；（3）不同材料对附着动物的定殖效果存在显著差异，钢材的附着动物生物量显著高于混凝土，平均是其8.3倍（P<0.05）。结果表明，在同一礁体单元内，混凝土更有利于大型藻类建植，钢材则更有利于附着动物定殖，二者组合使用不仅能够同步促进大型藻类和附着动物定殖的双效提升，从而实现了材料的功能互补，还可显著提高附着生物的生物量与多样性水平，为人工藻礁建礁材料筛选提供了新思路。
- 人工藻礁 /
- 材料筛选 /
- 大型藻类 /
- 群落演替 /
- 附着效果
Abstract: To address the limited habitat heterogeneity and suboptimal biological attachment capacity of single-material artificial algal reefs in seaweed bed restoration, this study designed a steel-concrete composite artificial algal reef. Twelve replicate reefs were fabricated and deployed in the Mashanli sea area of Rongcheng in August 2021. Attached macroalgae and animals colonizing both materials, as well as key environmental parameters at the reef site, were monitored in November 2021, June 2022, September 2022, May 2023, and August 2023. Results revealed: (1) the two materials differed significant in their macroalgal establishment effects, with the biomass of attached macroalgal on concrete being significantly higher than that attached on steel, reaching 1.7 times that on steel during the entire monitoring period on average (P < 0.05); (2) macroalgal community structure was jointly shaped by seasonality and reef age, progressing from an early Chlorophyta-dominated phase to a late Phaeophyta-dominated phase. Rhodophyta appeared transiently but never established a persistent dominant stage. After 20 months, a stable, top-level community dominated by Undaria pinnatifida emerged (Y = 0.99); (3) the colonization dynamics of attached animals varied significantly between two substrate materials. Specifically the biomass of attached animals on steel substrates was 8.3 times higher than that on concrete substrates (P < 0.05). These results indicate that, within identical reef units, concrete supported greater macroalgal establishment, whereas steel facilitated significantly higher settlement of attached animals. Combining both materials in artificial reef construction yields synergistic benefits: it concurrently promotes macroalgal development and attached animal colonization, thereby achieving functional complementarity, and significantly enhances both the total biomass and taxonomic diversity of sessile organisms. This dual-substrate strategy provides an evidence-based framework for optimizing material selection in artificial algal reef engineering.
- Artificial algal reef /
- Material selection /
- Macroalgae /
- Community succession /
- Attachment effect

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图 1 实验区域

Fig. 1 The experimental area

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图 2 人工藻礁立体结构示意图（A）与内部结构俯视图（B）

注：①为水体交换孔（Φ80 mm）；②为钢制立柱；③为半圆形钢板构件（Φ200 mm）；④为1/4圆弧无孔钢板底座；⑤为不同材料水平板。

Fig. 2 Schematic diagram (A) and internal structural perspective (B) of the artificial algal reef

Note: ① a water exchange aperture (diameter: 80 mm); ② a load-bearing structural steel column; ③ a semi-circular steel plate component (diameter: 200 mm); ④ a quarter-circular, non-porous steel base plate; and ⑤ a horizontally oriented composite plate composed of different materials.

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图 3 不同材料大型附着藻类生物量的变化

注：误差棒上小写字母表示相同材料不同时间之间存在显著差异，*表示相同时间不同材料之间存在显著差异（P<0.05）。

Fig. 3 Temporal changes in macroalgal biomass across substrate materials

Note: Lowercase letters on error bars denote significant differences among sampling times within each substrate material; and asterisks (*) denote significant differences between substrate types at each sampling time (P<0.05).

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图 4 不同材料藻类附着状况的变化

Fig. 4 Temporal changes in macroalgal settlement density across substrate types

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图 5 不同材料大型藻类香农-威纳多样性指数(A)、物种丰富度指数(B)和均匀度指数(C)的变化

注：S: 钢材; C: 混凝土; 误差棒上小写字母表示相同材料不同时间之间存在显著差异，*表示相同时间不同材料之间存在显著差异（P<0.05）。

Fig. 5 Temporal changes in Shannon-Wiener index (A), Margalef's index (B), and Pielou's index (C) of macroalgae between substrate types

Note: S: Steel; C: Concrete. Lowercase letters on error bars denote significant differences among sampling times within each substrate material; and asterisks (*) denote significant differences between substrate types at each sampling time (P<0.05).

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图 6 大型藻类群落聚类分析（A）与非度量多维尺度排序（nMDS）（B）

注：分组表示为“时间-材料”。S：钢材，C：混凝土。

Fig. 6 Cluster analysis (A) and non-metric multidimensional scaling (nMDS) ordination (B) of macroalgal assemblages across substrate types

Note: Samples are grouped by sampling time and substrate type (S: steel; C: concrete), with substrate as the primary fixed factor in multivariate analyses.

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图 7 不同材料附着动物生物量（A）与密度（B）的变化

注：S: 钢材 Steel; C: 混凝土 Concrete; 误差棒上小写字母表示相同材料不同时间之间存在显著差异，*表示相同时间不同材料之间存在显著差异（P<0.05）。

Fig. 7 Temporal changes in attached animal biomass (A) and density (B) across substrate materials

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表 1 水体环境指标调查结果

Tab. 1 Investigation results of water environmental indicators

指标 Parameter	2021.11	2022.06	2022.09	2023.05	2023.08
温度T（℃）	15.87 ± 0.15^d	24.93 ± 0.32^a	21.53 ± 0.25^c	15.40 ± 0.30^d	23.67 ± 0.38^b
盐度Sal（‰）	31.97 ± 0.06^d	29.57 ± 0.64^a	29.70 ± 0.36^ab	30.23 ± 0.06^bc	30.40 ± 0.17^c
流速V（m·s⁻¹）	0.13 ± 0.01^ab	0.14 ± 0.02^ab	0.15 ± 0.01^b	0.13 ± 0.01^ab	0.12 ± 0.02^a
溶解氧DO（mg L⁻¹）	7.60 ± 0.01^c	7.43 ± 0.04^d	6.68 ± 0.08^e	12.56 ± 0.02^a	9.06 ± 0.09^b
pH值	8.47 ± 0.06^a	8.30 ± 0.10^a	8.47 ± 0.15^a	8.46 ± 0.04^a	7.31 ± 0.09^b
$ {{\rm {PO}}_4^{3-}} $-P（mg L⁻¹）	0.020 ± 0.001^a	0.012 ± 0.001^b	0.016 ± 0.001^a	0.003 ± 0.001^c	0.006 ± 0.002^c
无机氮DIN（mg L⁻¹）	0.35 ± 0.01^b	0.28 ± 0.03^c	0.34 ± 0.01^c	0.55 ± 0.07^a	0.40 ± 0.03^b
注：小写字母表示不同调查时间之间存在显著差异（P<0.05）。
Note: Lowercase letters indicate statistically significant differences across survey time (P<0.05).

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表 2 人工藻礁大型附着藻类名录

Tab. 2 Species composition of macroalgae associated with artificial algal reefs

门 Phylum	种类 Species	2021.11		2022.06		2022.09		2023.05		2023.08		温度性质 Temperature adaptation nature
门 Phylum	种类 Species	S	C	S	C	S	C	S	C	S	C	温度性质 Temperature adaptation nature
绿藻门 Chlorophyta	肠浒苔 Enteromorpha intestinalis	+						+				CT
	缘管浒苔 Enteromorpha linza	+	+					+	+			WT
	假根羽藻 Bryopsis corticulans	+	+									WT
	孔石莼 Ulva pertusa			+	+	+	+	+	+	+		WT
	刺松藻 Codium fragile									+		WT
	硬毛藻 Chaetomorpha antennina					+	+					WT
	线形硬毛藻 Chaetomorpha linum			+	+	+	+	+	+	+		WT
红藻门 Rhodophyta	小珊瑚藻 Corallina pilulifera	+	+									ST
	角叉菜 Chondrus ocellatus							+	+			WT
	多管藻 Polysiphonia urceolata								+			LB
	繁枝蜈蚣藻 Grateloupia ramosissima								+			WT
	龙须菜 Asparagus schoberioides										+	WT
褐藻门 Phaeophyta	水云 Ectocarpus siliculosus	+										CT
褐藻门 Phaeophyta	裙带菜Undaria pinnatifida							+	+	+	+	WT
注：S表示钢材，C表示混凝土，+表示采到藻类，WT表示暖温带种类，CT表示冷温带种类，ST表示亚热带种类，LB表示亚寒带种类。
Note: S = steel; C = concrete; “+” denotes presence of the corresponding macroalgal species; WT = warm temperate; CT = cold temperate; ST = sub tropical; LB = lower boreal.

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表 3 不同材料大型藻类优势种及优势度

Tab. 3 Dominant macroalgal species and their relative dominance across substrate types

时间 Time	优势种 Dominant species	优势度 Relative dominance
时间 Time	优势种 Dominant species	钢材 Steel	混凝土 Concrete
2021.11	假根羽藻 Bryopsis corticulans	0.49	0.50
	缘管浒苔 Enteromorpha linza	0.30	0.42
	小珊瑚藻 Corallina pilulifera	0.06	0.08
	水云 Ectocarpus siliculosus	0.15	—
2022.06	孔石莼 Ulva pertusa	1.00	1.00
2022.09	孔石莼 Ulva pertusa	0.99	0.89
2022.09	硬毛藻 Chaetomorpha antennina	—	0.10
2023.05	裙带菜 Undaria pinnatifida	0.99	0.99
2023.08	裙带菜 Undaria pinnatifida	0.57	0.97
	孔石莼 Ulva pertusa	0.42	—
	龙须菜 Asparagus schoberioides	—	0.03

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表 4 不同调查时间两种材料表面绿藻门、红藻门和褐藻门生物量占大型附着藻类总生物量的比例

Tab. 4 Proportions of Chlorophyta, Rhodophyta and Phaeophyta biomass in the total attached macroalgal biomass on steel and concrete substrates survey times

时间 Time	门 Phylum	占比 Proportion
时间 Time	门 Phylum	钢材 Steel	混凝土 Concrete
2021.11	绿藻门 Chlorophyta	64.80	92.18
	红藻门 Rhodophyta	6.07	7.82
	褐藻门 Phaeophyta	10.89	—
2022.06	绿藻门 Chlorophyta	100.00	100.00
	红藻门 Rhodophyta	—	—
	褐藻门 Phaeophyta	—	—
2022.09	绿藻门 Chlorophyta	100.00	100.00
	红藻门 Rhodophyta	—	—
	褐藻门 Phaeophyta	—	—
2023.05	绿藻门 Chlorophyta	1.31	0.20
	红藻门 Rhodophyta	0.00	0.29
	褐藻门 Phaeophyta	98.68	99.52
2023.08	绿藻门 Chlorophyta	42.52	—
	红藻门 Rhodophyta	—	3.00
	褐藻门 Phaeophyta	57.48	96.99
注：—表示未检出，0.00表示已检出但占比极低。
Note: “—” denotes absence of detection; “0.00” indicates presence at trace levels.

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表 5 人工藻礁附着动物名录

Tab. 5 Species composition of attached animals associated with artificial algal reefs

门 Phylum	种类 Species	2021.11		2022.06		2022.09		2023.05		2023.08		温度性质 Temperature adaptation nature
门 Phylum	种类 Species	S	C	S	C	S	C	S	C	S	C	温度性质 Temperature adaptation nature
软体动物门 Mollusca	长牡蛎 Crassostrea gigas	+		+	+	+		+	+	+	+	WT
	紫贻贝 Mytilus edulis			+	+	+						WT
	东方缝栖蛤 Hiatella orientalis			+	+						+	WT
	短滨螺 Littorina brevicula								+			WT
	红带织纹螺 Nassarius succinctus									+		WT
	秀丽织纹螺 Nassarius festivus									+		WT
	布尔小笔螺 Mitrella burchardi										+	WT
	丽小笔螺 Mitrella bella										+	WT
节肢动物门 Arthropoda	四齿矶蟹 Pugettia quadridens							+				WT
节肢动物门 Arthropoda	大蜾蠃蜚 Corophium major									+		WT
脊索动物门 Chordata	玻璃海鞘 Ciona intestinalis					+				+	+	WT
脊索动物门 Chordata	柄海鞘 Styela clava			+	+					+		WT
环节动物门 Annelida	独齿围沙蚕 Perinereis cultrifera	+	+									WT
	长双须虫 Eteone longa	+										WT
	多齿全刺沙蚕 Nectoneanthes multignatha			+				+				WT
	沙蚕 Nereis succinea			+						+		WT
	无疣齿吻沙蚕 Inermonephtys inermis									+		WT
	寡鳃齿吻沙蚕 Nephtys oligobranchia									+	+	WT
	多鳃齿吻沙蚕 Nephtys polybranchia									+		WT
	不倒翁虫 Sternaspis scutata									+		WT
	网纹哈鳞虫 Harmothoё dictyophora									+	+	WT
	长须沙蚕 Nereis longior									+	+	WT
	树蛰虫 Pista cristata									+		WT
	长锥虫 Haploscoloplos elongates									+		WT
	拟特须虫 Paralacydonia paradoxa										+	WT
注：S表示钢材，C表示混凝土，+表示采集到附着动物，WT表示暖温带种类。
Note: S = steel; C = concrete; “+” denotes presence of the corresponding attached animal species; WT = warm temperate.

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表 6 附着动物生物多样性指数

Tab. 6 Diversity indices of attached animal assemblages across substrate types

时间 Time	材料 Material	香农-威纳多样性指数H' Shannon-Wiener index	物种丰富度指数D Margalef’s index	均匀度指数J Pielou’s index
2021.11	钢材	0.81 ± 0.25*	0.24 ± 0.10*	0.97 ± 0.05
	混凝土	0.00 ± 0.00	0.00 ± 0.00	—
2022.06	钢材	1.42 ± 0.02*	0.51 ± 0.01*	0.89 ± 0.01
	混凝土	0.85 ± 0.16	0.31 ± 0.01	0.78 ± 0.15
2022.09	钢材	0.46 ± 0.40	0.13 ± 0.11	1.00 ± 0.00
	混凝土	—	—	—
2023.05	钢材	0.52 ± 0.04	0.16 ± 0.00	0.77 ± 0.04
	混凝土	0.21 ± 0.37	0.06 ± 0.10	0.87 ± 0.05*
2023.08	钢材	2.11 ± 0.15	1.13 ± 0.10*	0.93 ± 0.02
	混凝土	1.71 ± 0.20	0.75 ± 0.14	0.96 ± 0.03
注：“*”表示相同时间不同材料间具有显著差异（P<0.05）。2021年11月混凝土表面仅记录到1种附着动物，故H'和D为0，J以“—”表示；2022年9月混凝土表面未记录到附着动物，故相关指数均以“—”表示。
Note: “*” denote significant differences between substrate types at each sampling time(P<0.05). In November 2021, only one attached animal species was detected on the concrete substrate; consequently H' and D were both zero, whereas J is undefined and reported as “—”. In September 2022, no attached animals species were detected on the concrete substrate; and therefore, all three diversity indices are undefined and reported as “—”.

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