The Structure of Coral Communities at Hurghada in the Northern Red Sea

Abstract. The community structure of hard and soft corals, with an emphasis on hard corals, was determined by means of line-transects on 9 on-and off-shore reefs of different type and different wave exposure in the Northern Red Sea near Hurghada in Egypt. Coral communities were found to differentiate along a horizontal wind- and wave-exposure gradient. Exposed communities were dominated by Acropora species, sheltered communities by Porites species, and semi-exposed communities by Millepora species. Also, vertical within-reef zonations following a depth gradient were observed; these were unique for each exposure-determined community type. Average transect diversity was highest on semi-exposed reefs, lowest on sheltered reefs. Reef slopes were more diverse than other reef zones. The observed community structure was compared with data from the literature, and widely distributed, roughly comparable hard and soft coral communities were identified.     

The Skeleton Eroding Band Disease on Coral Reefs of Aqaba, Red Sea

Abstract.  The occurrence and distribution of the coral disease Skeleton Eroding Band (SEB) has been studied at the Jordanian coast in the Gulf of Aqaba, the northernmost extension of the Red Sea. The SEB syndrome is caused by the colonial, heterotrich ciliate Halofolliculina corallasia : an advancing front of these protists destroys polyps, coenosarc, and surface of the coenosteum. In recent years SEB was registered in reefs of Australia, Mauritius, and the Red Sea. Along the Jordanian coast, disease frequency at four study sites was investigated and afflicted coral genera and species were documented. The study sites were MSS (Marine Science Station)-north, MSS-south, the tourist area near the middle of the coast, and the industrial area adjacent to the Saudi Arabian border. Corals and infections were counted inside randomly placed 1 m² frames in reef flats and in 5, 10, and 15 m depths. In order of increasing affectedness of sample sites, MSS-south was the healthiest (12%, 4%, 4%, 16% of infected corals from shallow to deep), followed by MSS-north (19%, 28%, 24%, 24%), the tourist area (51%, 23%, 16%, 41%), and the industrial area (14%, 30%, 39%, 31%). By far the most frequently encountered coral species were Acropora spp. and Stylophora sp., while relative infection-rates were highest among Seriatopora sp. (75%), as well as Stylophora sp., Hydnophora sp., and Galaxea sp. (50% each). This was followed in steadily decreasing order from 44% to 5% by Pocillopora , Mycedium , Montipora , Echinopora , Acropora , Lobophyllia , Goniastrea , Millepora , Platygyra , Fungia , Favia , Porites , Goniopora , Favites , and Pavona . SEB was found to a depth of 30 m, but may occur even deeper.     

海洋酸化、暖化对两种鹿角珊瑚幼虫附着及幼体存活的影响

大气CO₂浓度持续升高导致海洋酸化和暖化影响着造礁石珊瑚和珊瑚礁生态系统。为探明造礁石珊瑚早期生活史对海洋酸化和暖化的生理学响应，本文研究了温度（约28°C, 约30°C）和pCO₂（约570 μatm, 约1 300 μatm）以及两者协同作用对简单鹿角珊瑚（Acropora austera）和中间鹿角珊瑚（A. intermedia）早期生活史的影响。实验结果表明，升温（+约2.5°C）和酸化（约1 300 μatm）对两种鹿角珊瑚幼虫的附着率和死亡率均无显著影响。酸化显著降低了简单鹿角珊瑚幼体存活率（25.87%），但并不显著影响中间鹿角珊瑚幼体的存活率；升温对两种鹿角珊瑚幼体存活率无显著影响。升温（+约2.5°C）、酸化（约1 300 μatm）对简单、中间鹿角珊瑚幼虫的存活和附着过程的影响较小，但是酸化对简单鹿角珊瑚幼体存活的影响高于暖化。本文结果表明，珊瑚补充过程对海洋酸化和暖化的响应可能具有种类特异性，气候变化将逐渐改变造礁石珊瑚的群落结构。     

Assessment of the effectiveness of natural coral fragmentation as a dispersal mechanism for coral reef‐boring sponges

Eastern Pacific reefs are mostly made up of interlocking coral branches of Pocillopora, which are easily broken by physical forces associated with heavy swells and winds. In this study we investigated the potential of these coral fragments to enable propagation of boring sponges. For this, we quantified the frequency of occurrence and diversity of boring sponges in fragments of corals recently trapped among the branches of live colonies, and later tested the hypothesis that these sponges colonize new branches of corals. Nearly 80% of the coral colonies investigated had coral fragments among their branches, and 69% of these coral fragments contained boring sponges (11 species), some of these sponges in reproduction (23% of them carried oocytes). To test whether sponges inhabiting coral fragments could colonize new branching corals we transplanted them to healthy branches, and to branches whose living tissue was mechanically eliminated to simulate damage produced by grazing and death after bleaching and other causes of coral tissue mortality. All the transplanted coral fragments cemented to each new colony by means of calcification, and of the three sponge species tested (Cliona vermifera, Cliona tropicalis and Thoosa mismalolli) only C. vermifera was able to colonize both new living branches (26.9%) and cleaned branches (65.5%). The apparent capability of C. vermifera to colonize by direct contact may be another key ability of this species to maintain high frequency of occurrence in Pacific coral reefs. However, although C. tropicalis and T. mismalolli were not able to colonize new coral substrata by direct contact, coral fragments have the potential to contribute to local persistence of these sponges and to their dispersal, both by asexual (fragments) and sexual means (transport of sexual products). The present findings may partly explain the current increase of excavating sponges on deteriorating reefs with a large availability of dead branching corals.     

Coral reefs of the Glorieuses Islands,western Indian Ocean

The benthic fauna on reefs around the Glorieuses Islands, a small and protected Indian Ocean archipelago northwest of Madagascar, was surveyed in November 2015, focusing particularly on the orders Alcyonacea (soft corals and gorgonians) and Scleractinia (stony corals). The species richness of both groups was rather low for a protected, relatively pristine environment in the region. Though certain soft ‘fugitive’ alcyonacean species were noticeably abundant, other soft corals were rare. Sediment-tolerant faviid corals were abundant among the scleractinians. The sediment around the reefs was fine-grained and white, and appeared to be derived from abundant green macroalgae Halimeda and coralline seaweeds. The abundance of these, in turn, might have been attributable to nutrient enrichment from guano deposited on one of the islands by migratory seabirds, causing the aforementioned anomalies in coral biodiversity and abundance.     

Coral Recruitment and Regeneration on a Maldivian Reef Four Years after the Coral Bleaching Event of 1998. Part 2: 2001–2002

Abstract.  This paper is the follow-up study to a quantitative inventory of stony corals on a Maldivian reef after the bleaching event of 1998 [Loch, K., W. Loch, H. Schuhmacher & W. R. See, 2002; P.S.Z.N.: Marine Ecology, 23 (3): 219–236] covering the years 2001 and 2002, thereby extending the observation period from 1999 to 2002. Data collection was performed as previously in transects on the reef flat, Acropora tables at the reef rim, Porites lobata blocks and one Diploastrea heliopora colony. While between 2000 and 2001 the transects showed a noticeable increase of living scleractinian colonies of 6.1 per m² of reef surface per year, this had slowed to 0.7 per m² per year by 2002. Likewise, the colonisation of dead Acropora tables had changed from a yearly increase of 14.4 colonies per m² of table surface between 2000 and 2001 to a decrease of 9.4 between 2001 and 2002. The main constituent of the regeneration in transects as well as on Acropora tables was Pavona (Agariciidae), whereas Pocilloporidae and Acroporidae were almost completely missing. A new immigrant was Tubastraea micranthus . Continuous growth of regenerates on Porites presented a columnar form, and the growth rate of D. heliopora was reduced between 2001 and 2002. The influence on reef ecology in terms of coral substrate, species differentiation, growth rate changes, interaction with other organisms and the overall prospect of the reef development on a longer timescale is outlined.     

海南西岛珊瑚群落变化及其影响因素 *

在2018年的海南三亚西岛珊瑚群落分布和水质环境因子调查中, 共记录到造礁石珊瑚68种, 多孔螅2种。基于珊瑚礁底质类型的非度量多维标度分析, 可将西岛站位划分为区域1(东-南侧)和区域2(西侧)。全岛活珊瑚覆盖率为13.6%, 区域1和区域2分别为4.2%和27.7%, 区域2珊瑚多样性高于区域1。区域1珊瑚礁处在显著退化状态, 而区域2相对健康, 但是优势种类单一。与历史资料对比分析, 东侧站位造礁石珊瑚覆盖率从2011年的近40%降至2018年不足4%, 退化显著, 而西侧站位珊瑚礁无明显退化; 丛生盔形珊瑚(Galaxea fascicularis)逐渐取代分枝状的鹿角珊瑚成为绝对优势种类; 珊瑚礁区浊度、溶解无机氮(dissolved inorganic nitrogen, DIN)和喇叭藻的δ¹⁵N均值分别为2.54NTU、9.63μmol·L^-1和5.09‰, 均处在较高水平。相关性分析表明, 环境因子的空间变化并不能解释珊瑚群落的空间分布。因此, 推测人类活动引起的极端扰动事件可能是导致西岛珊瑚群落空间分布和东侧站位珊瑚近几年显著退化的主要原因, 如三亚河清淤在三亚湾随意倾倒引起的浑浊事件。应加强对三亚湾清淤和西岛人类活动的管控, 减轻颗粒物沉积和营养盐富集的影响, 促进西岛珊瑚礁逐渐恢复。     

海南岛西北部海域珊瑚礁造礁石珊瑚种类组成与分布

调查了海南岛西北部海域造礁石珊瑚种类的多样性及分布.结果表明:共发现50种造礁石珊瑚和1种多孔螅.海南岛西北部造礁石珊瑚优势种为澄黄滨珊瑚(Porites lutea)和丛生盔形珊瑚(Galaxea fascicularis),与历史资料相比优势种发生明显变化;海南岛西北部海域造礁石珊瑚覆盖率急剧下降,是由自然因素和人为活动共同造成;海南岛西北部海域是北部湾、琼州海峡和南海的重要交汇点,可能是造礁石珊瑚幼虫扩散的重要通道,对华南沿岸造礁石珊瑚资源自然恢复起到重要作用.因此,未来的海洋管理应该加强对珊瑚礁的保护力度,开发利用与环境保护并重,这样才能有效保护海南岛西北部海域的珊瑚礁.     

永兴岛及七连屿造礁石珊瑚近10年变化分析

根据2007-2016年珊瑚礁监测数据，永兴岛及七连屿共调查到造礁石珊瑚11科113种，常见种有鹿角杯形珊瑚Pocillopora damicornis、疣状杯形珊瑚Pocillopora Verrucosa、粗野鹿角珊瑚Acropora humilis、澄黄滨珊瑚Porites lutea、丛生盔形珊瑚Galaxea fascicularis、梳状菊花珊瑚Goniastrea pectinata及锯齿刺星珊瑚Cyphastrea serailia等。研究区域内造礁石珊瑚种类近10年（2007-2016年）来下降明显，其中永兴岛从39种下降到18种，下降率为53.85%；北岛从23种下降到17种，下降率为26.09%；赵述岛从46种下降到12种，下降率为73.91%；西沙洲从51种下降到15种，下降率为70.59%。群体形态群落中皮壳状种类数最为稳定，分枝类下降了59.26%；叶片状下降了75.00%；圆盘状下降了87.50%；团块状下降了57.14%。通过对研究区域造礁石珊瑚种类组成及群落形态等方面的变化分析可见，人为活动、长棘海星、珊瑚病害、台风及全球升温等因素逐渐影响到西沙海域的珊瑚礁生态系统。     

鹿回头佳丽鹿角珊瑚卵母细胞发育的组织学研究

海南省三亚市鹿回头海域珊瑚礁生态系统退化严重,为了恢复珊瑚礁资源,引进国际上最新的珊瑚礁恢复策略--捕获珊瑚幼虫安放到珊瑚礁退化区域,该策略的重点是准确掌握珊瑚的排卵时间.实验通过组织学分析珊瑚卵母细胞发育情况从而判定珊瑚的排卵时间,而野外的珊瑚幼虫捕获实验也证实我们对排卵时间的判断.通过观测卵母细胞的发育情况,我们确定佳丽鹿角珊瑚Acropora pulchra的排卵时间为4月底5月初.同时也证实佳丽鹿角珊瑚共生的虫黄藻Zooxanthellae是后天获得的.将组织学分析应用到珊瑚礁的恢复中来,使得以后大量的捕获珊瑚幼虫,加快对珊瑚礁退化区域进行恢复成为可能.     

Coral communities and reefs from Guerrero,Southern Mexican Pacific

Corals in the Eastern Pacific extend south from the Gulf of California to Ecuador and oceanic Chile, and west from Colombia to Clipperton Atoll. Nevertheless, large stretches of the Mexican Pacific remain fundamentally unstudied. Therefore, to assess the current conditions of coral communities, a coastal fringe ～300 km long (17°40′ N, 101°39′ W to 16°46′ N, 99°49′ W) was surveyed within the Southern Mexican Pacific, between 2005 and 2009. Fifteen stony coral species were identified at 13 coral communities and six Pocillopora‐dominated fringing reefs, with Pocillopora verrucosa and Pocillopora damicornis the primary contributing taxa. Reef development was identified in embayments or behind rocks or islands that offered shelter from northern and northwestern winds. Observations of Pocillopora effusus, Pocillopora inflata, Porites lobata, Pavona clavus, and Pavona varians expanded the species known geographic ranges by several degrees of latitude, suggesting reef building fauna comprised a mixture of widespread and relatively rare Eastern Pacific corals. Results indicated greater live coral cover in the Ixtapa‐Zihuatanejo area (15–73%) than in the Acapulco localities, which had high algal dominance; the reefs in the latter region exhibited high erosion. Regional differences are likely the result of long‐standing anthropogenic pressures around Acapulco since 1950, when it became an important tourist destination. This paper is the first detailed report of ecologically stressed corals and coral reefs from the state of Guerrero on the Mexican Southern Pacific coast.     

Coral colonization by the encrusting excavating Caribbean sponge Cliona delitrix

The Caribbean sponge Cliona delitrix is among the strongest reef space competitors; it is able to overpower entire coral heads by undermining coral polyps. It has become abundant in reefs exposed to organic pollution, such as San Andrés Island, Colombia, SW Caribbean. Forty‐four sponge‐colonized coral colonies were followed‐up for 13 months to establish the circumstances and the speed at which this sponge advances laterally into live coral tissue and the coral tissue retreats. Cliona delitrix presence and abundance was recorded at seven stations to interpret current reef space and coral species colonization trends. The spread of C. delitrix on a coral colony was preceded by a band of dead coral a few millimeters to several centimeters wide. However, the sponge was directly responsible for coral death only when live coral tissue was within about 2 cm distance; coral death became sponge advance‐independent at greater distances, being indirectly dependent on other conditions that tend to accelerate its retreat. Cliona delitrix advanced fastest into recently killed clean coral calices; however, sponge spread slowed down when these became colonized by algae. The lateral advance of C. delitrix was slower than other Cliona spp. encrusting excavating sponges, probably owing to the greater depth of its excavation into the substratum. Cliona delitrix prefers elevated portions of massive corals, apparently settling on recently dead areas. It currently inhabits 6–9% of colonies in reefs bordering San Andrés. It was found more frequently in Siderastrea siderea (the most abundant local massive coral), which is apparently more susceptible to tissue mortality than other corals. Current massive coral mortality caused by C. delitrix could initially change the relative proportions of coral species and in the long‐term favor foliose and branching corals.     

海南昌江沿岸海域石珊瑚的物种多样性及其分布

2009年3月,采用水下照相及摄像方法,对海南昌江沿岸海域的石珊瑚物种多样性、分布及覆盖度进行了调查.结合历史资料,共发现石珊瑚10科15属28种和1未定种.其中鹿角珊瑚属（Acropora）、角蜂巢珊瑚属（Favites）、叶状珊瑚属（Lobophyllia）等为优势类群,常见分布珊瑚种类有角孔珊瑚（Gonioporasp.）、标准蜂巢珊瑚（Favia speciosa）、五边角蜂巢珊瑚（Favites pentagona）、多孔鹿角珊瑚（Acropora millepora）、十字牡丹珊瑚（Pavona decussata）、伞房叶状珊瑚（Lobophyllia corymbosa）等.石珊瑚主要分布于海尾镇三联村以南海域,覆盖率最高为12%.本次调查在海南昌江沿岸海域记录的珊瑚种类数多于以前的记录,但少于海南南岸的三亚鹿回头,也少于北部的广西涠洲岛、广东徐闻、香港等海域,随着调查次数的增加,预计会有更多的种类被发现.该海域是北部湾珊瑚幼虫交流的中间过渡区,对珊瑚礁的保护与恢复具有重要意义.     

中国珊瑚礁资源状况及其药用研究调查 Ⅰ.珊瑚礁资源与生态功能

对中国珊瑚礁生物物种组成、资源状况、分布特征及其生态功能进行了调查和评价.中国珊瑚礁主要分布在中国南海诸岛海域和唬南、广东、广西、福建、台湾等近岸海域.中国造礁石珊瑚已记录21科,56属295种,对珊瑚礁生态系统结构与生态功能的维持具有重要作用.八放珊瑚亚纲中柳珊瑚已记录7科,28属,75种,软珊瑚有6科,23属,113种,是现代海洋药物研究的热点生物资源.中国珊瑚礁具有造礁与消波护岸、维持珊瑚礁生态系统和生物多样性、提供生物资源和材料、生物地球化学与科学研究、文化教育与生态旅游等多种生态功能与价值.     

Change in coral reef benthic communities in the Lembeh Strait and Likupang, North Sulawesi, Indonesia

Anthropogenic impacts and natural disturbances have been intense recently in the global scale, affecting the composition of coral reef benthic communities from coral to algal dominated reefs. However, this condition does not always occur considering corals are able to recover when the stressors falter. This study aims to investigate the change in coral reef benthic communities and the relationship among benthic categories. The study was carried out in 2014 and 2016 at five sites, three sites in the Lembeh Strait and two sites in Likupang, North Sulawesi Province. Underwater Photo Transect(UPT) was used at depth of around 4–6 m in slope areas. The result indicated that the benthic communities were slightly changing: the percent covers of hard corals, sponges, soft corals, macroalgae and substrate categories were not significantly different between the years but category of others, particularly seasonally growing hydroid, increased significantly, occupying the available substrates and overtopping other benthos surrounding. The study also found that there was a significant relationship between the change in benthic gradient and the number of hard coral colonies: when the composition becomes less complex, the number of colony declines. In contrast, the hard coral diversity remained unchanged, suggesting the coral reefs apparently have an ecological resilience(sustainable species diversity) against the change although ecological complexity declines. In addition, the hard coral cover was significantly correlated with soft coral and sponge covers, which did not change significantly among the years. In general, the coral reefs in North Sulawesi might experience a temporary blip due to the increasing percent cover of others, and be predicted to recover as there was no indication of soft corals and sponges to increase significantly. However, it is necessary to investigate the dynamic of benthic communities in different depth gradients to gain a comprehensive understanding as the communities respond differently to the light intensity.     

Coral bleaching and habitat effects on colonisation of reef fish assemblages: An experimental study

Degradation and mortality of corals is increasing worldwide and is expected to have significant effects on coral reef fish; hence studies on these effects are essential. In the present study, a field experiment was set up within Mafia Island Marine Park in Tanzania (East Africa) to examine the effects of bleaching and habitat structure on colonisation of coral reef fish assemblages. Live and bleached staghorn coral Acropora formosa was transplanted onto plots in a site dominated by sand and rubble, and the experimental design comprised of three treatments: live coral, bleached coral and eroded coral rubble. There was an immediate increase (within 24 h) in fish abundance and diversity in the two treatments with standing corals. Overall, live and bleached coral plots showed similar effects, but differed from the eroded coral plots which had a much lower abundance and diversity of fish. In general, fish species diversity changed with time over the study period while fish abundance did not. Multivariate analyses showed that while there were differences in fish assemblage structure between standing corals and the eroded coral treatment, there was neither a difference between live and bleached coral treatments nor any temporal effects on fish assemblage structure. Our findings suggest that physical structure and complexity of habitat have stronger effects on colonisation of reef fish assemblages than changes in coral health (such as bleaching) which do not affect coral structure. This may have important implications for appropriate coral reef management.     

西沙赵述岛海域珊瑚礁生态修复效果的初步评估

珊瑚恢复是一个协助退化、受损的珊瑚礁生态系统恢复的过程.近10 a来,西沙珊瑚礁生态系统不断退化,至2009年珊瑚覆盖率已下降至不足10%.针对西沙群岛珊瑚礁生态系统的退化状况,2011年12月于西沙赵述岛海域投放人工礁基并开展珊瑚移植实验,比较了人工修复区和自然恢复区1a后的修复效果.结果显示,投放礁基且进行珊瑚移植的区域,移植珊瑚的存活率不到10%;投放礁基但未进行珊瑚移植的区域,珊瑚补充量高达6~7 ind/m2;自然恢复区其珊瑚覆盖率提高了19.4%.依据珊瑚礁生态修复效果评估方法,人工修复且进行移植珊瑚的区域的生物修复权重值为27.5,人工修复但未移植珊瑚区及自然恢复区的生物修复权重值为39.3,珊瑚修复生物指标为亚健康.我们的结果表明,在健康、有良好珊瑚幼虫补充的珊瑚礁生态系统中,珊瑚礁群落可以自行恢复,其恢复只是一个时间的问题;而在缺少珊瑚幼虫附着基底的区域,通过人为增加一些附着基底,可加速其恢复进程.     

DARWIN'S Coral Reef Research - a Review and Tribute

Abstract. In rememberance of the death of CHARLES DARWIN 100 years ago, a review is given on his coral reef research from the point of view of present knowledge. Although during DARWIN'S time no one was aware of the changing level of the glacial sea or of the important function of the symbiotic zooxanthellae in the life of hermatypic corals, his observations and considerations on the ecology of corals and on the formation of coral reefs are relevant to this day. But of course his ideas need new causal interpretations. The origin of the different reef types can no longer be explained by subsidence alone, but by different modes biologists and geologists described during the following century. To summarize all the many existing theories, the all enclosing "theory of compensation" is put forward here in honour of this great man.     

Coral Reef Community Structure at Caño Island, Pacific Costa Rica

Abstract. Around the Biological Reserve of Caño Island, Pacific Costa Rica, there are five large coral reef flats (with size ranges of 0.8–4.2 ha) built mainly of dead Pocillopora spp. At present, they are covered mainly by crustose coralline algae and microatolls of Porites lobata . From the upper reef slope to the reef base several corals grow in small patches (e. g., Pavona clavus, Pavona varians, Pavona gigantea, Gardineroseris planulata, Psammocora superficialis, Pocillopora elegans, Pocillopora damicornis); the massive coral Porites Iobata is predominant. Pocilloporid species are predominant on most other eastern Pacific reefs.
The Caño Island reef is typical of a community whose structure has been controlled by both physical (in shallow water) and biological (in deeper water) factors. Shallow reef areas are influenced by strong wave action and extreme low tides. The distribution, abundance, and feeding preferences of corallivorous organisms (e. g., Acanthaster planci, Arothron meleagris, Pseudobalistes naufragium, Quoyula monodonta) on the deeper reef suggest that most pocilloporids are affected and limited by them. Although there is no evidence of any predator on Porites lobata at Caño Island, the triggerfish Pseudobalistes naufragium breaks off fragments of the coral while searching for food. These fragments often survive to form new colonies. Together, this dispersal mechanism, rapid injury recovery, and high resistance to environmental stress seem to enhance the distribution and dominance of the massive coral Porites lobata at Caño Island.     

Patterns of Coral Distribution and Benthic Space Partitioning on the Fringing Reefs of Southern Taiwan

Abstract. The distribution pattern of corals and benthic space partitioning patterns on the fringing reefs of southern Taiwan were studied by the line-transect method. The bathymetric distribution of corals in the study area was basically homogeneous except on the protected reef slopes and the exposed reef fronts and terraces. Corals on protected reef slopes are mainly foliaceous, including Montipora foliosa, Merulina ampliata, Mycedium elephantotus , and Pachyseris speciosa , but the dominant species on a reef slope vary from site to site. The formation of the foliaceous coral community is possibly related to a complex of environmental factors, chance, and biological interactions. The reef fronts and submarine terraces of exposed areas are dominated by alcyonacean corals. The most abundant species are Sarcophyton trocheliophorum, S. crassocaule , and Lobophytum sarcophytoides ; their distributions are relatively homogeneous. The formation of alcyonacean-dominated assemblages can be related to the special adaptive strategies of these corals. In regard to space partitioning patterns, the major space occupiers on those reefs exposed to storm disturbances are alcyonacean soft corals, scleractinian corals, and algae; on reefs protected from storms, scleractinian corals and algae dominate. Two conspicuous features of the study area are the high proportion of space occupied by algae and the abundant unoccupied space. The dense algal cover is possibly caused by sewage pollution and overfishing of the reef area. The unoccupied space is most likely related to the high frequency of typhoon disturbances.