臺灣博碩士論文加值系統

小琉球為台灣唯一的珊瑚礁島嶼，曾為台灣地區珊瑚生長最繁茂，海洋生物多樣性最高的區域之一。近年來，小琉球珊瑚礁生態系遭受天然及人為壓力，導致珊瑚覆蓋率持續減少。本研究欲了解 2019 至 2020 年小琉球周圍海域珊瑚群聚狀態及珊瑚礁年間變化，於小琉球漁業資源保育區內設置四個樣點(漁福、厚石、杉福、花瓶岩)，參考國際珊瑚礁總體檢調查法於每個樣點 5 m 及 10 m 水深佈設長 50 m 的穿越線（transect），採用樣點擷取法(point intercept method )，記錄底質覆蓋率；此外，於穿越線兩側 5 m 處擺放 50*50 cm 的樣框，調查珊瑚種類、生長型態、珊瑚生態功能群 (rK-S)及分析其生物多樣性指數，使用聚類分析(Cluster Analysis)、多元尺度分析(MDS)探討珊瑚種類組成的變異程度，同時配合相似性分析(ANOSIM)判讀小琉球珊瑚是否有分群現象，再使用相似度貢獻分析法(SIMPER) 從中找出平均差異度前 10%的珊瑚種類，進而探討小琉球珊瑚群聚的時空變化；為了更深入探討小琉球珊瑚群聚可能的影響因子，於 2020 年加入 4 樣點 5m 水深的水質及沉積物粒徑調查，並以主成份分析(PCA)探討小琉球海域影響珊瑚的主要環境因子。研究結果顯示，二年間的底質類型皆以岩石(RK)為主(平均值 37.20±12.12%)；2020 年硬珊瑚(HC)有略為增加 (平均值24.73±14.30%)；值得注意的是 2020 年藻類(NIA)明顯增加(平均值 24.60±13.02%)，超過硬珊瑚數量(平均值 24.08±14.24%)，可能顯示有「相變」的潛在性。珊瑚覆蓋率結果兩年皆屬於「劣化」程度(2019年平均值22.61±10.56%；2020年平均值24.73±14.30%)，且珊瑚生長型態皆以耐壓型為多(團塊形及表覆形)；相似性分析結果顯示，兩年珊瑚群聚相似性分析具顯著差異(R=0.97，P<0.05)，相似度貢獻平均差異高達 75.63%，珊瑚種類貢獻度前六名為火珊瑚(Millepora sp.)、鐘形微孔珊瑚(Porites lutea)、團塊微孔珊瑚(Porites lobata)、羅氏盤星珊瑚(Dipsastraea rotumana)、實心角菊珊瑚(Favites halicora)及鱗指形軟珊瑚(Sinularia scabra)；造成此差異可能與水質條件的氨氮(NH4-N)及總有機碳(TOC)；沉積物條件的黏土、泥及極粗砂有關，推測小琉球海域受自然環境及人為擾動的影響較高。綜合而論，小琉球海域可能受自然環境變遷及人為擾動影響，導致珊瑚礁生態的生存受到壓迫而持續衰敗，建議以漁福及厚石（含水深 5m 及10m)劃設重點保育示範區，作為小琉球珊瑚群聚的避護所。另外，應立即進行遊客人數總量管制，降低人為活動所造成的環境影響，以利小琉球珊瑚生態系的永續發展。

Hsiao-Liu-Chiu is the only one island formed by coral reefs in Taiwan with the highest diversity of marine organisms. In recent years, Hsiao-Liu-Chiu has historically been experienced multiple natural and anthropogenic disturbances, resulting in a sharp reduction of coral coverage. In order to reveal the states of coral communities of Hsiao-Liu-Chiu, four sampling locations including Yufu, Houshi, Shanfu, and Huapingyan in the Fishery Conservation Zone were surveyed from 2019 to 2020. At 5m and 10m water depth of each location, coverage of benthic assemblages was surveyed using point intercept method with 50 m long line transect, and coral growth forms, colony number, and species composition in 500 m2 were surveyed using a photo-transect method with 30 photographs (50 x 50 cm). The cluster analysis and multidimensional scaling (MDS) ordination of coral communities was conducted to reveal the grouping of surveyed sites. Analysis of similarity (ANOSIM) was applied to separate different groupings from MDS and the Similarity Percentage (SIMPER) analysis was used to reveal the species that contributed to top 10% species in cumulative similarities within groups with a multivariate analysis package Primer 6.0. The r-K-S analysis was used to indicate the conservation class of each location. In order to understand the potential factors contributed to the change of coral communities of Hsiao-Liu-Chiu, water quality and sedimentary size at 5m water depth of each location were measured and analyzed using Principal Component Analysis (PCA). Our results shown that Rock (RK) was the most common substrate occupied (average 37.20±12.12%) in this 2-year period; Hard coral (HC) (24.73±14.30%) slightly increased in 2022. It should be noted that Algae (NIA) with 24.60±13.02% was higher than HC (24.08±14.24%) in 2020, suggesting the possibility of phase shift. The coral coverage in these
two years were in the deterioration state, ranging from 22.61±10.56% to 24.73±14.30% and dominated by the stress-tolerated corals with massive and encrusting growth form. The results
from ANOSIM indicated that there was a significantly different between coral communities in
2019 and 2020 (R=0.97，P<0.05) and the average dissimilarity was 75.63%. The top six dominant species contributing to this difference were Millepora sp., Porites lutea, Porites lobata, Dipsastraea rotumana, Favites halicora, Sinularia scabra. The NH4-N, Total organic carbon (TOC), clay, silt, and coarse sand could be the factors related to the change of coral communities, suggesting Hsiao-Liu-Chiu is under high level of natural and anthropogenic disturbances. Herein,We suggest that the Yufu and Houshi at 5 m and 10 m water depth should be the refuge for corals in the future and thus the coral communities in situ need to be protected. At the same time, total number of tourists should also be controlled for the sustainable development of the coral ecosystem in Hsiao-Liu-Chiu.

壹、 前言 1

一、珊瑚礁的重要性 1
二、珊瑚礁現況及影響珊瑚礁可能的因子 2
三、小琉球的地理位置、氣候環境及海域概況 3
四、影響小琉球珊瑚群聚的因子 4
五、文獻回顧 5
六、研究目的 8

貳、 材料及方法 9

一、調查時間及地點 9
二、調查方法 9

(一)底質類型及珊瑚覆蓋率調查 9
(二)珊瑚種類和豐富度調查 9
(三)珊瑚生長形態調查及辨別 10
(四)環境因子調查 10
(五)相關資料蒐集 10

三、統計分析 11

(一)多樣性指數分析 11
(二)聚類分析(Cluster Analysis) 12
(三)多元尺度分析(Multidimensional Scaling) 12
(四)相似度貢獻分析法(SIMPER) 12
(五)主成份分析(Principal Component Analysis) 12
(六)珊瑚生態功能群分析(r-K-S) 13
(七)統計軟體運用 13

參、 結果 14

一、底質類型及珊瑚覆蓋率調查結果 14

(一)漁福樣點 14
(二)厚石樣點 15
(三)杉福樣點 16
(四)花瓶岩樣點 16
二、珊瑚種類多樣性調查結果 17

(一)漁福樣點 17
(二)厚石樣點 18
(三)杉福樣點 18
(四)花瓶岩樣點 19
(五)珊瑚群聚組成分析結果 19
(六)珊瑚群聚組功能群分析 20
三、環境因子調查結果 21

(一)水質 21
(二)沉積物粒徑 21

肆、 討論 22

伍、 參考文獻 28

陸、 附件 54

交通部中央氣象局 (2022) https://www.cwb.gov.tw/V8/C/

交通部觀光局統計資料庫 (2020) https://stat.taiwan.net.tw/scenicSpot

何旻杰 (2014)。台灣北部鹽寮灣海域珊瑚群聚之研究。國立臺灣大學海洋研究所，博 士論文

林幸助、邵廣昭、黃守忠 (2020)。我國海洋生態調查監測網與監測規範建立之整體規劃。

國家海洋研究院委託之專題研究成果報告。國家海洋研究院。

邵廣昭、呂學榮、鄭有容、陳天任 林綉美、吳書平、林嘉瑋(2019)。臺灣南部海域人工魚礁區、水產動植物繁殖保育區生態調查。行政院農業委員會 108 年度國土生態保育綠色網絡建置計畫。

屏東縣琉球鄉公所 (2022) https://www.pthg.gov.tw/liuchiu/Default.aspx

珊瑚礁總體檢報告 (2009)。環境資訊協會，https://www.slideshare.net/reefcheck/2009-2728328

珊瑚礁總體檢報告 (2015)。環境資訊協會，https://www.slideshare.net/reefcheck/2015-56326719

珊瑚礁總體檢報告 (2016)。環境資訊協會，https://www.slideshare.net/reefcheck/2016-70458046

珊瑚礁總體檢報告 (2017)。環境資訊協會，https://www.slideshare.net/reefcheck/2017-108449450

珊瑚礁總體檢報告 (2018)。環境資訊協會，https://www.slideshare.net/reefcheck/2018-146707832

海洋委員會海洋保育署 (2022)https://www.oca.gov.tw/ch/index.jsp

張水楷、劉莉蓮、樊同雲、廖德裕 (2019)。屏東縣海洋及漁業事務管理所委託服務工作計畫。

陳正平、王豐寓、劉名允、何詩琥 (2017)。珊瑚礁樂園--小琉球，科學發展月刊(563)，36-43。

陳昭倫、郭兆揚、黃雅怡、鍾愛琪、陳彥嘉、陳亭儒、蕭安孜、謝孟芸、蔡政翰 (2019)。108 年珊瑚礁生態系調查計畫成果報告書。海洋委員會海洋保育署委託報告

楊東霖（2012）。小琉球海域珊瑚礁底棲群聚結構之時空變化。國立東華大學海洋生物多樣性及演化研究所，碩士論文。

楊萬全 (1997)。高屏溪流域和屏東平原的水資源，師大地理研究報告(27)，81-104。

廖德裕 (2021)。示範海域海洋生物資訊蒐集及資源調查-南部海域。國家海洋研究院委託之專題研究成果報告。

戴昌鳳 (2007)。台灣海域珊瑚礁分布及珊瑚多樣性調查研究。行政院農委會漁業署 96年度科技計畫研究報告。

戴昌鳳 (2022). 台灣珊瑚全圖鑑(下)：軟珊瑚。台灣：貓頭鷹出版社。

戴昌鳳、洪聖雯 (2009)。台灣珊瑚礁圖鑑。台灣：貓頭鷹出版社。

戴昌鳳、秦啟翔 (2019) 墾丁國家公園八放珊瑚生態圖鑑。墾丁國家公園管理處。

戴昌鳳、秦啟翔、鄭安怡 (2013)。東沙珊瑚生態圖鑑。台灣：內政部營建署海洋國家公園管理處。

戴昌鳳、鄭有容 (2020). 台灣珊瑚全圖鑑(上)：石珊瑚。台灣：貓頭鷹出版社。

羅柳墀 (2013)。遊憩衝擊對小琉球潮間帶生物多樣性與族群空間分布的影響. 地理學報(69), 25-46。

Abidin SZZ, Mohamed B (2014) A review of SCUBA diving impacts and implication for coral reefs conservation and tourism management SHS Web of Conferences. EDP Sciences, pp01093

Adam TC, Burkepile DE, Holbrook SJ, Carpenter RC, Claudet J, Loiseau C, Thiault L, Brooks AJ, Washburn L, Schmitt RJ (2021) Landscape‐scale patterns of nutrient enrichment in a coral reef ecosystem: implications for coral to algae phase shifts. Ecological Applications
31: e2227

Anthony KR, A. Kleypas J, Gattuso JP (2011) Coral reefs modify their seawater carbon chemistry–implications for impacts of ocean acidification. Global Change Biology 17:3655-3666

Beck MW, Losada IJ, Menéndez P, Reguero BG, Díaz-Simal P, Fernández F (2018) The global flood protection savings provided by coral reefs. Nature communications 9: 1-9

Birkeland C (2015) Coral reefs in the Anthropocene Coral reefs in the Anthropocene. Springer, pp 1-15

Bozec Y-M, O'Farrell S, Bruggemann JH, Luckhurst BE, Mumby PJ (2016) Tradeoffs between fisheries harvest and the resilience of coral reefs. Proceedings of the National Academy of Sciences 113: 4536-4541

Burke L, Reytar K, Spalding M, Perry A (2011) Reefs at risk revisited. World Resources Institute

Carpenter KE, Abrar M, Aeby G, Aronson RB, Banks S, Bruckner A, Chiriboga A, Cortés J, Delbeek JC, DeVantier L (2008) One-third of reef-building corals face elevated extinction risk from climate change and local impacts. Science 321: 560-563

Cheng Y-R, Chin C-H, Lin D-F, Wang C-K (2020) The probability of an unrecoverable coral community in Dongsha Atoll Marine National Park due to recurrent disturbances. Sustainability 12: 9052

Clarke K, Gorley R (2005) PRIMER: Getting started with v6. PRIMER-E Ltd: Plymouth, UK 931: 932

Clarke K, Green R (1988) Statistical design and analysis for a'biological effects' study. Marine Ecology Progress Series: 213-226

Connell JH (1978) Diversity in tropical rain forests and coral reefs: high diversity of trees and corals is maintained only in a nonequilibrium state. Science 199: 1302-1310

Costanza R, De Groot R, Sutton P, Van der Ploeg S, Anderson SJ, Kubiszewski I, Farber S, Turner RK (2014) Changes in the global value of ecosystem services. Global environmental change 26: 152-158

Cramer KL, Donovan MK, Jackson JB, Greenstein BJ, Korpanty CA, Cook GM, Pandolfi JM (2021) The transformation of Caribbean coral communities since humans. Ecology and Evolution 11: 10098-10118

Dai C (2005) Dong-sha Atoll in the South China Sea: Past, present and future Islands of the world VIII international conference, Kinmen Island, Taiwan

De K, Nanajkar M, Mote S, Ingole B (2020) Coral damage by recreational diving activities in a Marine Protected Area of India: Unaccountability leading to ‘tragedy of the not so commons’. Marine Pollution Bulletin 155: 111190

Di Franco A, Milazzo M, Baiata P, Tomasello A, Chemello R (2009) Scuba diver behaviour and its effects on the biota of a Mediterranean marine protected area. Environmental Conservation 36: 32-40

Duignan M (2019) ‘Overtourism’? Understanding and managing urban tourism growth beyond perceptions: Cambridge case study: Strategies and tactics to tackle overtourism. United Nations World Tourism Organisation (UNWTO), pp 34-39

Dyshlovoy SA, Honecker F (2020) Marine compounds and cancer: The first two decades of XXI century. Multidisciplinary Digital Publishing Institute, pp 20

Edinger EN, Risk MJ (2000) Reef classification by coral morphology predicts coral reef conservation value. Biological Conservation 92: 1-13

Fabricius KE (2011) Factors determining the resilience of coral reefs to eutrophication: a review
and conceptual model. Coral reefs: an ecosystem in transition: 493-505
Ferrario F, Beck MW, Storlazzi CD, Micheli F, Shepard CC, Airoldi L (2014) The effectiveness of coral reefs for coastal hazard risk reduction and adaptation. Nature communications 5:1-9

Gratwicke B, Speight MR (2005) Effects of habitat complexity on Caribbean marine fish assemblages. Marine Ecology Progress Series 292: 301-310

Hardiman N, Burgin S (2010) Recreational impacts on the fauna of Australian coastal marine ecosystems. Journal of environmental management 91: 2096-2108

Hawkins JP, Roberts CM (1992) Effects of recreational SCUBA diving on fore-reef slope communities of coral reefs. Biological Conservation 62: 171-178

Hodgson G (1999) A global assessment of human effects on coral reefs. Marine Pollution Bulletin 38: 345-355

Hodgson G (2000) Coral reef monitoring and management using Reef Check. Integrated coastal zone management 1: 169-179

Hoegh-Guldberg O, Eakin C, Hodgson G, Sale P, Veron J (2015) Climate change threatens the survival of coral reefs. International Society for Reef Studies (ISRS) 2015: 4

Hoegh-Guldberg O, Mumby PJ, Hooten AJ, Steneck RS, Greenfield P, Gomez E, Harvell CD, Sale PF, Edwards AJ, Caldeira K (2007) Coral reefs under rapid climate change and ocean acidification. science 318: 1737-1742

Huang Y-C, Hsieh H, Huang S-C, Meng P-J, Chen Y-S, Keshavmurthy S, Nozawa Y, Chen C (2011) Nutrient enrichment caused by marine cage culture and its influence on subtropical coral communities in turbid waters. Marine Ecology Progress Series 423: 83-93

Johnson GW, Ehrlich R (2002) State of the art report on multivariate chemometric methods in environmental forensics. Environmental forensics 3: 59-79

Karcher DB, Roth F, Carvalho S, El-Khaled YC, Tilstra A, Kürten B, Struck U, Jones BH, Wild C (2020) Nitrogen eutrophication particularly promotes turf algae in coral reefs of the central Red Sea. PeerJ 8: e8737

Kleypas JA, McManus JW, Meñez LA (1999) Environmental limits to coral reef development: where do we draw the line? American zoologist 39: 146-159

Kornder NA, Riegl BM, Figueiredo J (2018) Thresholds and drivers of coral calcification responses to climate change. Global change biology 24: 5084-5095

Kuo C, Ho M, Chen Y, Yang S, Huang Y, Hsieh H, Jeng M, Chen C (2018) Status of Coral Reefs in East Asian Seas Region: Taiwan. Ministry of the Environment of Japan: 33-40

Li X-b, Huang H, Lian J-s, Liu S, Huang L-m, Yang J-h (2013) Spatial and temporal variations in sediment accumulation and their impacts on coral communities in the Sanya Coral Reef Reserve, Hainan, China. Deep Sea Research Part II: Topical Studies in Oceanography 96: 88-96

Luckhurst B, Luckhurst K (1978) Analysis of the influence of ubstrate variables on coral reef fish communities. Marine Biology 49: 317-323
McCauley DJ, Pinsky ML, Palumbi SR, Estes JA, Joyce FH, Warner RR (2015) Marine defaunation: animal loss in the global ocean. Science 347: 1255641

McCook LJ (1999) Macroalgae, nutrients and phase shifts on coral reefs: scientific issues and management consequences for the Great Barrier Reef. Coral reefs 18: 357-367

Moberg F, Folke C (1999) Ecological goods and services of coral reef ecosystems. Ecological economics 29: 215-233

Molinski TF, Dalisay DS, Lievens SL, Saludes JP (2009) Drug development from marine natural products. Nature reviews Drug discovery 8: 69-85

Morrison TH, Hughes TP, Adger WN, Brown K, Barnett J, Lemos MC (2019) Save reefs to rescue all ecosystems. Nature Publishing Group

Newman SP, Meesters EH, Dryden CS, Williams SM, Sanchez C, Mumby PJ, Polunin NV (2015) Reef flattening effects on total richness and species responses in the C aribbean. Journal of Animal Ecology 84: 1678-1689

NOAA, Coral Reef Watch(2022)。https://coralreefwatch.noaa.gov/

Perez III K, Rodgers KS, Jokiel PL, Lager CV, Lager DJ (2014) Effects of terrigenous sediment on settlement and survival of the reef coral Pocillopora damicornis. PeerJ 2: e387

Puotinen M, Drost E, Lowe R, Depczynski M, Radford B, Heyward A, Gilmour J (2020) Towards modelling the future risk of cyclone wave damage to the world's coral reefs.Global Change Biology 26: 4302-4315

Randazzo-Eisemann Á , Garza-Pérez JR, Penié-Rodriguez I, Figueroa-Zavala B (2021) 25 years of multiple stressors driving the coral-algae phase shift in Akumal, Mexico. Ocean & Coastal Management 214: 105917

Reef Check Foundation(2022)。https://www.reefcheck.org/

Shokri MR, Mohammadi M (2021) Effects of recreational SCUBA diving on coral reefs with an emphasis on tourism suitability index and carrying capacity of reefs in Kish Island, the northern Persian Gulf. Regional Studies in Marine Science 45: 101813

Sobel J, Dahlgren C (2004) Marine reserves: a guide to science, design, and use. Island Press

Spalding M, Burke L, Wood SA, Ashpole J, Hutchison J, Zu Ermgassen P (2017) Mapping the global value and distribution of coral reef tourism. Marine Policy 82: 104-113

Spalding M, Spalding MD, Ravilious C, Green EP (2001) World atlas of coral reefs. Univ of California Press

Teh LS, Teh LC, Sumaila UR (2013) A global estimate of the number of coral reef fishers. PLoS One 8: e65397

Terrón-Sigler A, León-Muez D, Peñalver-Duque P, Torre FE (2016) The effects of SCUBA diving on the endemic Mediterranean coral Astroides calycularis. Ocean & Coastal Management 122: 1-8

Veron JEN (1986) Corals of Australia and the Indo-Pacific.University of Hawaii Press, Honolulu

Wilson SK, Babcock RC, Fisher R, Holmes TH, Moore JA, Thomson DP (2012) Relative and combined effects of habitat and fishing on reef fish communities across a limited fishing gradient at Ningaloo. Marine Environmental Research 81: 1-11

WoRMS Editorial Board。(2022)。https://www.marinespecies.org/

Yang R, Chi K, Hu S, Chen H (1975) Corals, fishes and benthic biota of Hsiao-Liu-Chiu. Institute of Oceanography, National Taiwan University, Special publication 7, pp 53

Zakai D, Chadwick-Furman NE (2002) Impacts of intensive recreational diving on reef corals at Eilat, northern Red Sea. Biological Conservation 105: 179-187

Zaneveld JR, Burkepile DE, Shantz AA, Pritchard CE, McMinds R, Payet JP, Welsh R, Correa A, Lemoine NP, Rosales S (2016) Overfishing and nutrient pollution interact with temperature to disrupt coral reefs down to microbial scales. Nature communications 7: 1-12