Macroalgae in the coral reefs of Eilat (Gulf of Aqaba, Red Sea) as a possible indicator of reef degradation
Introduction
Coral reefs have deteriorated extensively during the past decades as a result of environmental disturbances of both anthropogenic and natural sources (Graham et al., 2006, Rogers and Miller, 2006, Wilkinson, 2004). According to Wilkinson (2004) 20% of the world’s coral reefs have been effectively destroyed and approximately 24% of the remaining reefs in the world are under imminent risk of collapse through human pressure. One way of rehabilitating damaged reefs is via reef management. A major impediment to management and prevention of further deterioration of the reefs lies in our inability to predict such decline by identifying the early warning signs of potentially dramatic community changes. Scientists are still unable to fully recognize the conditions that lead to the collapse of reef communities, even on reefs that are relatively well studied (Bellwood et al., 2004). For example, the coral reefs of the Caribbean have been thoroughly studied in relation to various human and natural effects, such as overfishing, hurricanes, sea urchin disease outbreak, and bleaching events (Hughes, 1994, Ostrander et al., 2000), providing quantitative data regarding reef degradation (Hughes et al., 1999). Although such features of reef degradation as loss of macro-fauna, reduced fish stocks, a shift from fish-dominated to echinoid-dominated herbivory, and reduced coral recruitment, have been well documented, the trajectory to ecosystem collapse remained unrecognized. Therefore, the ecosystem’s failure to regenerate remained unforeseen (Bellwood et al., 2004).
Degraded reefs often experience a transition in the structure of their benthic communities from dominant reef-building corals to overtaking marine macroalgae, a phenomenon termed ‘phase shift’ (Done, 1992, Hughes, 1994). Such shift may have significant ecological implications among them loss of structural complexity (Done, 1999, Spencer and Viles, 2002) that can affect coral recruitment (McCook et al., 2001), fish recruitment, competition, and predation (Öhman et al., 1998). These changes may eventually lead to a decline in species richness and biodiversity and, therefore, to an unstable ecosystem (Graham et al., 2006, Kiessling, 2005). Interactions among the coral reef occupants have been demonstrated in various models, in which stony corals, algae, ‘bottom-up’ (i.e. eutrophication) and ‘top-down’ (i.e. presence of grazers) factors are considered to be the main forces controlling the coral reef equilibrium (Done, 1992, Knowlton, 1992, Mumby, 2006). According to these models, the balance of scleractinian corals and algae is not expected to be interrupted when driven by nutrient enrichment alone. However, the combined effects of reduced herbivory together with enhanced nutrient supply (eutrophication) can shift the equilibrium in favor of algal dominancy (McCook, 1999, Miller, 1998, Thacker et al., 2001).
The coral reefs of the Red Sea are considered to be among the “healthiest” in the world (Hodgson, 1999), or, at least, in “relatively good health” (Wilkinson, 2004). Defining coral reefs based on their state of health, especially when they have been classified as being in a “good condition”, can lead to a subsequent lack of concern by decision-makers and, consequently, no action being taken when a situation arises that requires countermeasures. A small-scale evaluation of the status of local reefs along the Red Sea can thus be beneficial in providing a tool to obtain a reliable focal assessment for any given reef.
The coral reefs of Eilat (northern tip of the Gulf of Aqaba, Red Sea) are among the most intensively studied, spatially-limited, coral reefs in the world. Despite the intensive research, the data related to the reef state is relatively poor, and therefore researchers have conflicting opinions regarding the reef’s health. These conflicts are well expressed in two contradictory reports by an International Expert Team (IET), which aimed at providing the Israeli Government with a reliable synopsis of the northern Gulf of Eilat’s ecological state. According to the first IET report “The unique coral reef in Eilat has been deteriorating at an alarming rate over the last three decades…the Israeli coral reefs apparently suffer from lost diversity, decrease in coral cover, low rates of coral–larval settlement and recruitment” (Atkinson et al., 2001). However, in the second report, the IET states that “there was more evidence presented of reef improvement than of reef decline” (Atkinson et al., 2004). The problem in determining the reef state was further reflected in other publications, which also demonstrate contradictory conclusions regarding the reef state (Ben-Tzvi et al., 2004, Fishelson, 1995, Loya, 2007, Rinkevich, 2005), and the potential factors responsible for reef decline (Abelson et al., 2005, Glassom et al., 2005, Loya et al., 2004). Most of the degradation-related research that was carried out in the coral reefs of Eilat was focused on changes in several reef community indices associated with corals while neglecting the accompanied changes in benthic algae and their potential role which have often been overlooked.
The present study was aimed at examining the structure and dynamics of benthic-algal assemblages, notably turf algae, as a tool to indicate a reef’s state of health. Turf algae dominancy may have significant ecological implications for coral reefs, including its adverse effects on recruits (Birrell et al., 2005). Moreover, turf algae may compete for space with coralline algae, which are known for their essential role in coral reefs in inducing larval settlement and metamorphosis (Heyward and Negri, 1999, Kitamura et al., 2007), and for their significant contribution to reef construction by calcification (Fabricius and De’ath, 2001). Therefore, examination of the spatial distribution and effects of turf algae in the coral reefs of Eilat is expected to shed light on the actual state of health of the reefs.
In view of the fact that algae spread and occupy vacant substrates that were formerly occupied by reef-building corals and coralline algae (Jompa and McCook, 2003, Miller, 1998, Miller and Hay, 1998), the assessment of changes in algal assemblages can function as an additional index that may contribute to filling in the gap in the accumulated data regarding the state of health of Eilat’s reefs.
Section snippets
Study sites
The study was carried out at six sites along the northern tip of the Gulf of Aqaba (Eilat, Israel); three sites along the coast of Eilat and three sites along the coast of Aqaba, Jordan (Fig. 1). The Eilat sites were surveyed for a period of 12 months (from July 2005 to June 2006) for examination of coral–algal dynamics at 5 and 10 m depth. The study sites were along the southern coast of Eilat: at the northern part of the Nature Reserve (NR), at the Inter-University marine Institute (IUI) and at
Algal definition
Algal turf cover was defined as an array of several thin filamentous species forming a gray homogeneous layer (Fig. 2). Species in the algal mosaic are represented by minuscule strings (up to a few mm) which trap large amounts of sediment in their web. All organisms were identified to order level and two dominant species were identified to the genus level (Table 1). The most dominant species of the algal mosaic was Sphacelaria sp. (Phaeophyta), while all the others belonged to the same order
Discussion
Algal take-over is a common phenomenon in degraded reefs and the most dominant species in the turf complex studied here was Sphacelaria sp. (Phaeophyta). This same species was reported also in the Great Barrier Reef (GBR, Australia) as a main component in algal take-over. A homogenous layer of Sphacelaria sp. and other red alga species (Rhodophyta) was observed to occupy large areas of bleached corals shortly after the mass bleaching event in 1998 (Diaz-Pulido and McCook, 2002). In contrast,
Acknowledgments
This study was funded by the Red Sea Marine Peace Park (RSMPP) program of the US agency for international development – Middle Eastern Regional Cooperation (USAID-MERC) to A.A. The authors thank Dr. Athanasios Athanasiadis from Goteborg University for his help in algal identification, and Ms. N. Paz for her editorial assistance. We also thank Mr. S. Martinez for his assistance in data collection and the directors and staff of the Interuniversity Institute (IUI) in Eilat and the Marine Science
References (54)
- et al.
Coral recruitment to the reefs of Eilat, Red Sea: temporal and spatial variation, and possible effects of anthropogenic disturbances
Marine Pollution Bulletin
(2005) - et al.
Mass transport from pollution sources to remote coral reefs in Eilat (Gulf of Aqaba, Red Sea)
Marine Pollution Bulletin
(1999) - et al.
Deterioration Index (DI): a suggested criterion for assessing the health of coral communities
Marine Pollution Bulletin
(2004) - et al.
Simulation of wind-driven circulation in the Gulf of Elat (Aqaba)
Journal of Marine Systems
(2000) - et al.
Effects of algal turfs and sediment on coral settlement
Marine Pollution Bulletin
(2005) - et al.
Methods and interpretation of coral recruitment studies: effects of anthropogenic disturbance?
Marine Pollution Bulletin
(2005) A global assessment of human effects on coral reefs
Marine Pollution Bulletin
(1999)Recruitment of coralline crusts and filamentous turf algae in the Galapagos Archipelago – effect of simulated scour, erosion and accretion
Journal of Experimental Marine Biology and Ecology
(1991)- et al.
Characterization of a natural inducer of coral larval metamorphosis
Journal of Experimental Marine Biology and Ecology
(2007) - et al.
Larval development and settlement behavior of the gorgonian coral Plexaura kuna (Lasker, Kim and Coffroth)
Journal of Experimental Marine Biology and Ecology
(1996)
How to influence environmental decision makers? The case of Eilat (Red Sea) coral reefs
Journal of Experimental Marine Biology and Ecology
Nutrient enrichment caused by in situ fish farms at Eilat, Red Sea is detrimental to coral reproduction
Marine Pollution Bulletin
Settlement strategies and distribution patterns of coral-reef fishes
Journal of Experimental Marine Biology and Ecology
What do we know about Eilat (Red Sea) reef degradation? A critical examination of the published literature
Journal of Experimental Marine Biology and Ecology
Bioconstruction, bioerosion and disturbance on tropical coasts: coral reefs and rocky limestone shores
Geomorphology
Experimental ecology of rocky intertidal habitats: what are we learning?
Journal of Experimental Marine Biology and Ecology
Confronting the coral reef crisis
Nature
Reproductive cycle and developmental processes during embryogenesis of Clavularia hamra (Cnidaria, Octocorallia)
Acta Zoologica
Space partitioning by stony corals soft corals and benthic algae on the coral reefs of the northern Gulf of Eilat (Red Sea)
Helgolander wiss. Meeresunters
Calcification by crustose coralline algae on the northern Great Barrier Reef, Australia
Limnology and Oceanography
Seasonality in algal assemblages on upwelling-influenced coral reefs in the Colombian Caribbean
Botanica Marina
The fate of bleached corals: patterns and dynamics of algal recruitment
Marine Ecology Progress Series
Phase shifts in coral reef communities and their ecological significance
Hydrobiologia
Coral community adaptability to environmental change at the scales of regions, reefs and reef zones
American Zoologist
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