Changes of scleractinian coral communities in the Daya Bay in the last 40 years
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摘要: 本研究采用文献调研、现场调查和室内模拟试验等方法对大亚湾造礁石珊瑚的种群结构演变及其成因进行了探讨。2020年7月现场调查共发现造礁石珊瑚39种，块状的扁脑（Platygyra）、蜂巢（Favia）和角蜂巢（Favites）等是湾内优势珊瑚。1983/1984−2020年，大亚湾珊瑚群落演变的主要特征是曾经优势的鹿角珊瑚（Acropora）和蔷薇珊瑚（Montipora）从湾内基本消失。综合全球珊瑚礁退化大背景和大亚湾独特的水环境分析发现：大亚湾珊瑚群落退化是多方面因素共同作用的结果，其中珊瑚本身的脆弱性和环境中持续的夏季异常高温与低溶解氧含量的共同作用可能是湾内霜鹿角珊瑚（A. pruinosa）退化的主要原因。从湾内珊瑚群落恢复的种源考量，建议加强对刷洲东岛珊瑚群落的保护。Abstract: In order to disclose the changes of coral communities in the Daya Bay, literature analysis, field investigation and laboratory simulating experiments were performed. A total of 39 scleractinian coral species were detected in the 2020 field investigation, and massive coral Platygyra, Favia and Favites were dominant in the bay. The inlet coral community change was featured by the near completely lost of once dominant coral Acropora and Montipora during 1983/1984−2020. Analysis based on the background of global coral degradation and the specialty of the seawater environment in the Daya Bay indicated that the coral community degradation in the Daya Bay is a result of multiple stressors, and the susceptibility of Acropora and the frequent occurrence of high temperature and low content oxygen may be the main cause of inlet A. pruinosa lost. At last, as a potential larva source of inlet coral restoration, coral community of Shuazhou East Island should be protected seriously.
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Key words:
- Daya Bay /
- scleractinian coral /
- community change /
- high temperature /
- low dissolved oxygen
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图 2 2020−2021年夏季大亚湾珊瑚群落调查站位
Fig. 2 Coral community investigation sites in summers of 2020−2021 in the Daya Bay
图 3 珊瑚模拟试验装置示意图
自然降氧试验条件：陈海水，放入氧气控制装置，关闭通气，自然光，室温；低盐度胁迫试验：培养体系为低盐度的海水，通气，自然光，室温；沉积物悬浮胁迫试验：陈海水，加入沉积物，自然通气，自然光，室温
Fig. 3 Schematic diagram of the coral stress experiment set-up
Natural oxygen reduction test condition: old seawater, add the oxygen control set-up, close ventilation, natural light, room temperature; low salinity stress test: low salinity seawater, ventilation, natural light, room temperature; sediments suspension stress test: old seawater, add sediment, natural ventilation, natural light, room temperature
图 5 2021年07月29日至8月1日大亚湾温度、盐度、溶解氧含量和叶绿素含量垂直分布
SM. 三门岛；D. 大辣甲；X. 小辣甲；B. 白沙洲；S. 南部；S2. 南部第二站位；N. 北部；图例后面的数字代表CTD记录号
Fig. 5 The vertical distribution of seawater temperature, salinity, dissolved oxygen content and chlorophyll content of theDaya Bay from July 29 to August 1, 2021
SM. Sanmen Island; D. Dalajia; X. Xiaolajia; B. Baishazhou; S. south; S2. south second station; N. north; the number after the legend represents the CTD record number
图 6 大亚湾鹿角珊瑚分布在相对较深的位置
a. 大辣甲刷洲东岛北面；b. 三门岛小横洲北面；c−f. 大辣甲西侧；d−f. 水深略浅，其珊瑚颜色较其他站位要淡很多；红色箭头指示鹿角珊瑚；a−c、f显示鹿角珊瑚生长位置相对较深
Fig. 6 The relative low lying niches of Acropora in the Daya Bay
a. North of East Island of Shuazhou, Dalajia; b. north of Xiaohengzhou, Sanmen Island; c−f. west of Dalajia; d−f. the seawater is low, and the coral color is a little pale; the red arrows indicated Acropoa; a−c, f. indicated coral live in relative low lying niches
图 7 大亚湾珊瑚的生物侵蚀
a, b. 刺冠海胆对块状珊瑚的侵蚀；c, d. 钻孔动物对滨珊瑚的侵蚀；e, f. 草皮海藻对块状珊瑚的侵蚀
Fig. 7 Bioerosion of coral in the Daya Bay
a, b. Bioerosion of Diadema setosum to mass coral; c, d. bioerosion of burrowing animals to Porites; e, f. bioerosion of turf algae to mass coral
图 8 高温低溶解氧含量胁迫下鹿角珊瑚和滨珊瑚的病症变化
a−c. 鹿角珊瑚在31°C和3 mg/L溶解氧作用下白化症状随时间的变化。a. 试验起始阶段；b. 60 h时远离生长点部位开始白化，箭头指示起始白化位点；c. 69 h时整株珊瑚白化，组织脱落，死亡。d−f. 滨珊瑚在32°C和4 mg/L溶解氧作用下白化症状随时间变化。d. 试验起始阶段；e. 胁迫70 h，箭头指示珊瑚斑块状白化；f. 胁迫70 h后解除胁迫，珊瑚慢慢恢复了生长，该图为6个月后拍摄
Fig. 8 The symptom change of Acropora and Porites under high temperature and low dissolved oxygen content
a−c. Acropora under 31°C and 3 mg/L dissolved oxygen. a. The start of the experiment; b. polyps far from growing point began bleaching after 60 h of stressindicated the initial bleaching site; c. the whole coral branch bleached, tissue lost and dead after 69 h of stress. d−f. Temporal symptoms of Porites under 32°C and 4 mg/L dissolved oxygen. d. The start of the experiment; e. patchy bleaching happened after 70 h of stress, arrows indicated different patches; f. stress was removed after 70 h, then Porites restored its growth, photo was took 6 month after the stress
图 10 小辣甲滨珊瑚骨骼（采自4 m水深）年最低Sr/Ca和遮浪站年最高月均表层海水温度（多年平均温度28.2°C）距平（数据源于文献[24]）
Fig. 10 Departure of the annual minimum Sr/Ca of Porites skeleton collected from 4 m depth of the Xiaolajia Island, and the annual maximum monthly average sea surface temperature (the multi-year average is 28.2°C) recorded in Zhelang Station (the date are frm reference [24])
A1 2020年7月大亚湾珊瑚群落调查单样线（50 m）最高活珊瑚覆盖度（%）
A1 The maximum live coral cover (%) in a single 50 m transect during the investigation of July 2020
序号 珊瑚种类 白沙洲 小辣甲 大辣甲 三门岛 1 翼形蔷薇珊瑚Montipora peltiformis (Bernard, 1897) − 0.2 1.6 8.8 2 平展蔷薇珊瑚Montipora solanderi (Bernard,1897) − − − 6.4 3 霜鹿角珊瑚Acropora pruinosa (Brook, 1893) − + + 0.4 4 指形鹿角珊瑚Acropora digitifera (Dana, 1846) − − + 0.6 5 单独鹿角珊瑚Acropora solitaryensis (Veron and Wallace, 1984) − − + 3.2 6 罩柱群珊瑚Stylocoeniella guentheri (Bassett-Smith, 1890) − 0.4 − − 7 丛生盔形珊瑚Galaxea fascicularis (Linnaeus, 1767) − 0.4 − 0.2 8 浅薄沙珊瑚Psammocora superficialis (Gardiner, 1898) − 2.4 0.2 − 9 毗邻沙珊瑚Psammocora contigua (Esper, 1794) − 0.2 − − 10 十字牡丹珊瑚Pavona decussata (Dana, 1846) − 0.2 0.2 0.2 11 腐蚀刺柄珊瑚Hydnophora exesa (Pallas, 1766) 0.2 − − 0.2 12 盾形陀螺珊瑚Turbinaria peltata (Esper, 1794) − − 2.6 − 13 罗图马蜂巢珊瑚Favia rotumana (Gardiner, 1899) 0.2 0.8 1.2 1.4 14 标准蜂巢珊瑚Favia speciosa (Dana, 1846) 0.2 0.6 1.4 1.6 15 美龙氏蜂巢珊瑚Favia veroni (Moll and Borel-Best, 1984) − 0.2 1 − 16 圆纹蜂巢珊瑚Favia pallida (Dana, 1846) − − − 0.4 17 海洋蜂巢珊瑚Favia maritima (Nemenzo, 1971) − − − 0.4 18 秘密角蜂巢珊瑚Favites abdita (Ellis and Solander, 1786) 0.6 2.2 0.4 1.4 19 多弯角蜂巢珊瑚Favites flexuosa (Dana, 1846) 0.2 0.4 0.8 − 20 五边角蜂巢珊瑚Favites pentagona (Esper, 1794) 0.2 3.2 1 2.2 21 板叶角蜂巢珊瑚Favites complanata (Ehrenberg, 1834) − 0.2 0.6 − 22 海孔角蜂巢珊瑚Favites halicora (Ehrenberg, 1834) 0.2 1.4 2.4 1.2 23 罗素角蜂巢珊瑚Favites russelli (Wells, 1954) − − 0.2 − 24 粗糙菊花珊瑚Goniastrea aspera (Verrill, 1905) − 0.2 1.2 − 25 简短圆菊珊瑚Montastrea annuligera (Milne Edwards & Haime, 1849) − − − 0.4 26 肉质扁脑珊瑚Platygyra carnosus (Veron, 2000) 0.6 5 3.6 0.6 27 中华扁脑珊瑚Platygyra sinensis (Milne Edwards & Haime, 1849) 0.4 − 0.2 − 28 精巧扁脑珊瑚Platygyra daedalea (Ellis and Solander, 1786) 0.2 0.4 − − 29 小扁脑珊瑚Platygyra pini (Chevalier, 1975) − − 0.2 − 30 多孔同星珊瑚Plesiastrea versipora (Lamarck, 1816) 0.2 2.8 0.8 5 31 卷曲黑星珊瑚Oulastrea crispate (Lamarck, 1816) 0.4 − − − 32 白斑小星珊瑚Leptastrea pruinosa (Crossland, 1952) − − 0.2 1 33 紫小星珊瑚Leptastrea purpurea (Dana, 1846) − − − 0.6 34 锯齿刺星珊瑚Cyphastrea serailia (Forskal, 1775) − 0.6 − 0.6 35 澄黄滨珊瑚Porites lutea (Milne Edwards and Harime, 1851) 1.2 1.8 2 21.6 36 团块滨珊瑚 Porites lobata (Dana, 1846） − − − 0.6 37 柱角孔珊瑚Goniopora columna (Dana, 1846) − − 0.2 − 38 斯氏角孔珊瑚Goniopora stutchburyi (Wells, 1955) 0.8 − − 0.2 39 二异角孔珊瑚Goniopora duofasciata (Thiel, 1932) − − 0.4 − 注： “+”表示2021年调查有发现相应种类珊瑚；根据文献图片，粤东海区大陆认定的精巧扁脑和香港认定的肉质扁脑可能是同一种珊瑚；“−”代表在调查过程中未发现该种。 
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A2 大亚湾湾内活珊瑚覆盖度历年调查数据
A2 Records of live coral cover in the Daya Bay
调查年份 活珊瑚覆盖度/% 参考文献 大辣甲 小辣甲 白沙洲 1984/1985 72.7 78.1 77.7 张元林和邹仁林[31] 1991 32 32 − 温伟英等[32] 2003 22.9 58.8† − 林昭进等[30] 2006 20.5 − − 陈天然等[33] 2007 20.9 15.7 6.2 陈天然等[6] 2008 20.3 15.2 5.8 陈天然等[6] 2015 46.6‡ − − 张涛锂和练健生[29] 2020 9 20.2¶ − 郭峰等[28] 2020 11 7 2 本研究 注：†小辣甲北部小范围高值；‡采样站位不详，据前后数据分析可能也是小范围高值，据本文调查该值在大辣甲刷洲东岛是存在的；¶小辣甲东北部单站位，数据略高于本文调查；“−”代表无数据。
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