CAMBODIA MARINE CONSERVATION PROGRAMME Koh Rong Samloem, Cambodia

Transcription

1 CAMBODIA MARINE CONSERVATION PROGRAMME Koh Rong Samloem, Cambodia CMM Phase 142 Science Report 1 st April 31 st June

2 Staff Members Name Jenn Reitz (JR) Gen Labram (GL) Position Principle Investigator Assistant Research Officer 2

3 Table of Contents 1. Introduction Training Briefing Science lectures Field work training Research Work Program Survey areas Fish surveys Introduction Methodology Results Discussion Coral cover and other substrates surveys Introduction Methodology Results Discussion Benthic invertebrate surveys Introduction Methodology Results Discussion Proposed work program for next phase References

4 1. Introduction The Cambodian coastline, stretching 435 km along the Gulf of Thailand, is an especially productive marine ecosystem owing to the influx of nutrients from five major river systems reducing the salinity of the marine habitat (Touch, 1995). Thus far, 474 different species of fish from 105 different families have been identified, as well as mammals (dolphins, dugongs), reptiles (sea snakes, sea turtles) cephalopods (squid, octopus), crustaceans (crabs, shrimp, lobsters), molluscs (snails, sea slugs, bivalves), cnidarians (jellyfish, coral, sea anemones) and echinoderms (star fish, sea urchins, sea cucumbers) among other species which have not yet been recorded (Killeen, 2012). Cambodia has 69 coastal islands. Coral reefs, sea grass beds and mangroves fringe many of these; all are ecosystems that provide critical habitats for countless marine species. Coral reefs are one of the most diverse ecosystems on the planet. Southeast Asia contains some of the most species-rich reefs on earth and is home to the Coral Triangle; considered the epicenter of global marine biodiversity (Carpenter and Springer, 2005). The main historical threats to coral reefs were storms and volcanic eruptions. More recently however, anthropogenic pressures such as coastal development, overfishing and use of destructive fishing methods, ocean acidification and pollution have proven to be incredibly detrimental to the health of coral reefs. More than half of Southeast Asia s reefs are classified as high risk. This has primarily been caused by coastal development and fishing related pressures (Chou et al., 2002). Increases in coastal tourism can have a detrimental impact on coastal ecology and reef ecosystems if not properly managed (Hawkins and Roberts, 1994). Unregulated development can lead to substantial and potentially irreversible environmental degradation; through the construction of resorts and associated transportation infrastructure, overuse of water resources, increased fuel consumption and dumping of sewage and litter (Davenport and Davenport, 2006). Cambodia s minister of tourism, Thong Khon, reports that tourism rates in Cambodia are increasing. In 2013, 4.2 million foreign tourists visited Cambodia, an increase of 17.5% from the previous year (Cambodian Ministry of Tourism, 2013). Published information about the current state of Cambodia s reef systems is limited. Past studies have produced estimates of the number of species that may inhabit the reef ecosystem, including hard and soft corals, marine fish and molluscs. However, there are few studies of the reef s current condition, thus there is a need for additional studies to provide accurate data on the current status of these critical habitats (Chou et al., 2002; Wilkinson and Souter, 2008). 4

5 2. Training 2.1 Briefing Sessions Briefing sessions are conducted with research assistants (RAs) upon arrival at camp (Table 1). Table 1. Briefing sessions conducted during phase 142 Briefing session Introduction to CMM project Health and safety Medical briefing Camp life and duties Presenter JR JR JR JR 2.2 Science Lectures After the initial briefing session, RAs were given science lectures covering the background and rationale behind the Cambodia Beach Conservation project. Information relating to the different habitats and species of Koh Rong Samloem, as well as current conservation issues in Cambodia was also provided through a series of lectures (Table 2). The initial lectures included topics such as awareness of coastal hazards, the types of marine habitats in Cambodia, the forms and function of the coral reef habitat and reef fish morphology. Research assistants then proceeded to learn the reef fish that are surveyed using a series of flash cards. After successfully learning the fish species, RAs moved on to learning benthic substrate, invertebrate species and survey methodology. A fifth lecture, Coral cover, other substrates and survey methodology was presented at this time, followed by learning the types and classifications of substrates via flash cards. The final component of the main training module was a lecture covering basic invertebrate types and identification, combined with survey methodology and accompanied by a series of flash cards. During phase 142, a lecture specific to coastal ecology and conservation in Cambodia was completed and added to the basic training modules. There is also a series of lectures available for periods of inclement weather when in-water work is not possible. These lectures include topics such as: mangroves, coral reefs and climate change, marine pollution and Marine Protected Areas (MPAs). 5

6 Table 2. Science lectures delivered during phase 142 Lecture Coastal Hazards Coastal Ecology and Conservation in Cambodia Coral Reefs - Form and Function Reef Fish Morphology Coral cover, other substrates and survey methodology Benthic invertebrates and survey methodology Presenter JR JR JR/GL JR/GL JR/GL JR/GL 2.3 Fieldwork Training After the first four science training lectures (Table 2), RAs underwent a series of reef fish fieldwork training tests before being allowed to survey. After passing a series of tests with flash cards, in-water species tests were conducted. In order to pass the test, the most frequently seen species had to be correctly identified three times each, before being allowed to survey. This test is administered on a oneto-one basis as attempts in earlier phases with larger groups proved difficult. Following completion of the in-water species identification test, an in-water size differentiation test was carried out. These tests ensure that the RAs are able to accurately estimate reef fish size. Following the training for reef fish surveys, benthic substrate composition and invertebrate identification training sessions were carried out using a similar procedure. Practice surveys were conducted on the beach and in-water to ensure RAs were fully familiar with the surveying methodology. During this time, RAs also received training on any equipment they were unfamiliar with, such as how to use a compass or transect line. Following successful completion of fieldwork training, RAs were ready to start collecting survey data. 6

7 3. Research Work Program 3.1 Survey areas CMM is located in M Pai Bai village on Koh Rong Samloem Island, which is situated approximately 23 km west of Sihnaoukville on the South Coast of Cambodia. There are currently two survey sites in the waters surrounding M Pai Bai (10 34 N E). House Reef (Fig. 1), located on the east side of the village, is a coral reef located within a proposed conservation area, currently in the planning stages (personal communication, Coral Cay Conservation). Sunset Reef (Fig. 2), on the west side of the village, is predominantly rocky with corals and other types of substrate scattered throughout. Between the two reefs, there are currently a total of twelve active transect sites. However, tidal patterns at Sunset have prohibited data collection for a large portion of phase 142. As a result, there is only sufficient data available from House Reef s transects for analysis this phase (Table 3). Table 3. Transect locations and bearings in use at House Reef on Koh Rong Samloem. Transect Reef Bearing ( ) Average depth phase 141 (m) Average depth phase 142 (m) Average depth over both phases (m) 1 House Reef House Reef House Reef 50 4 House Reef 50 5 House Reef House Reef

8 Figure 1. House Reef on the east side of M Pai Bai Village. Figure 2. Sunset Reef, located on the west side of M Pai Bai village. 8

9 3.2 Fish surveys Introduction Reef fish diversity and abundance are important for supporting the ecosystems and communities on the island of Koh Rong Samloem. Both are key components for maintaining proper ecosystem function and in addition to their ecological importance, provide both a source of animal protein for the local population and support a number of different livelihoods such as fishing, diving and tourism. Thus, the health of coral reefs in the area is integral to the local economy, as most stakeholders rely on them in some capacity as a source of income. Factors that may impact diversity and abundance include overfishing, storm damage, terrestrial development, increasing sea-surface temperature, ocean acidification and sea level rises (Cheal et al., 2002; Zhao et al., 2009). Spatial and temporal monitoring of reef fish will provide an indication of the impact these disturbances are having on the reef. Effective monitoring efforts must be long-term as certain species in the Pomacentridae and Chaetodontidae families can show a delayed response to disturbance (Lewis, 1998) Methodology Underwater visual census adapted from reef check methodology (Reef Check, 2007) was used for data collection. Transect sites were selected for surveying dependent upon visibility and tide levels. At each of the survey sites, the abundance and size of 63 select families and species of fish were recorded. These species have been selected based on ecological and commercial importance. Variables such as transect number, sea state, cloud cover (percentage), tidal level and start time were recorded on entry into the water. At the marker buoy of the specified transect, start depth was measured using a weighted measuring tape. One surveyor subsequently set off on a predefined bearing, swimming slowly on that heading whilst the second surveyor laid the tape out to a distance of 20 m. At 20 m, the team waited approximately two minutes and then swam back along the length of the transect with one surveying 2.5 m left of the tape and the other surveying 2.5 m right of the tape, both recording species abundance and size (Fig. 3). On returning to the start of the transect, the team waited two further minutes and swam back along the length of the transect, again recording species abundance and size as they went. This two-minute period allows fish to return to their usual positions along the reef and mitigates the disturbance caused by the surveyors themselves. At the end of the survey, one team member kept a visual reference of the transect end point whilst the other reeled the tape in and returned to the end point. The end point water depth was then measured and recorded; all data was then verified by field staff before entry into database. 9

10 Figure 3. Baseline Survey Protocol (BSP) procedure for a 20 m transect. If any fish are seen within the 25 m 2 box, they are recorded by the surveyor Results Since the project began data collection at its present location on Koh Rong Samloem, 135 fish abundance surveys have been carried out on House Reef and Sunset Rock Reef in total. Eighty-four surveys were completed during phase 141 and 51 surveys occurred during phase 142. Due to extremely low water levels over transect sites located on Sunset Rock Reef in the previous phase, there are insufficient amounts of data for analysis from that site. During phase 141, higher species diversity was observed during the surveys on House Reef compared to Sunset (Table 4). No fish species new to the area have been observed during surveys this phase. Table 4. Species observed during surveys on House Reef and at Sunset Reef. Fish Species House Reef Sunset Crescent Wrasse Blue Streaked Cleaner Wrasse Tripletail Wrasse Chequerboard Wrasse Freckled Grouper Bluelined Grouper 10

11 Chocolate Grouper Honeycomb Grouper Spotted Coral Grouper Longfin Grouper Baramundi Grouper Peacock Grouper Starry Grouper Java Rabbitfish Golden Rabbitfish Virgate Rabbitfish Indian Goatfish Freckled Goatfish Eight Banded Butterflyfish Long Beak Coral Fish Blue-spotted Ribbon Tail Ray Checked Snapper Other Snapper Spp. Damselfish Spp. Monacle Bream Spp. Cardinalfish Spp. Sweeper Spp. Parrotfish Spp. Fusilier Spp. Squirelfish Spp. 11

12 Mean ± SE abundance of most commonly observed reef fish found at House Reef CMM 142 Science Report Emperor Spp. Porcupine Spp. Puffer Spp House Reef As with the previous phase, the most frequently observed families on House Reef were Damselfish spp., Cardinalfish spp., and Butterflyfish spp. (Fig. 4). The mean abundance of Damselfish spp., Butterflyfish spp. and Cardinal spp. remained relatively constant across the six transects. The mean abundance of other species observed on House Reef was considerably lower (Fig. 5). Total mean abundance was fairly consistent across transects at House Reef, with no significant differences (Fig. 6). This is in contrast with results from the previous phase, which indicated that transect 4 had significantly lower total mean abundance (296.7 ± 34.8 SE) than transects 3, 5, and 6 (Fig. 7) Butterflyfish Cardinal fish Damselfish Fish family Figure 4. Mean ± SE abundance of most commonly observed reef fish found at House Reef 12

13 Mean ± SE abundance of reef fish found at each transect Mean ± SE abundance of other reef fish found at House Reef CMM 142 Science Report Fish species Figure 5. Mean ± SE abundance of other reef fish found at House Reef Transect Figure 6. Mean ± SE abundance of reef fish found at transects on House Reef in phase

14 Mean ± SE abundance of reef fish found at each transect CMM 142 Science Report Transect Figure 7. Mean ± SE abundance of reef fish found at transects on House Reef in phase Discussion The data collected during this phase show many similarities to the results of the fish abundance and diversity surveys conducted during the previous phase and in the project s former location on Koh Smach. Damselfish continue to be the most commonly observed species at survey sites on both Koh Smach and Koh Rong Samloem. This may be partially explained by the sheer number of species included within the Damselfish family. Other factors that may explain why they are observed with such frequency include the availability of suitable habitats, food resources, breeding conditions and their size. Damselfish spp. tend to be of a relatively small size and thus are less likely to be regarded as a suitable catch for local fishermen. A 2003 study by De Lopez identified mullet, mackerel, snapper and sardines as the main catch brought in by fishermen in the Ream National Park area, supporting this theory. High abundance of other small species, such as Cardinalfish and Butterflyfish (Fig. 5), provides further support. Both snapper and mullet spp. were surveyed at Koh Rong Samloem and their numbers were found to be low on both reefs. During the previous phase, the total abundance of reef fish (regardless of species) showed some variation between transects across the same reef (Fig. 7). This was initially attributed to changing habitats across the reef; earlier studies have demonstrated that reef fish assemblages can change significantly across the reef shelf (Williams, 1982) due to changes in habitat complexity and composition of the coral community (Friedlander and Parish, 1998; Lewis, 1998). However, the results from this phase indicate no significant differences between total mean abundance across transects on House Reef. The dissimilarities observed 14

15 between phases are likely to be due to sampling effect as research activity in the area is still relatively recent. Nevertheless, it is important to continually reassess whether there are significant differences in fish abundance and assemblages and the conservation implications of this. In small areas with varying fish assemblages, appropriate zone planning is often necessary (Jennings et al., 1994) Coral cover and other substrate surveys Introduction In coral reef ecosystems, declining coral cover and condition has been linked to declines in fish biodiversity (Jones et al., 2004; Wilson et al., 2006). Thus, it is important to survey coral and other substrates to ascertain the overall health of a reef. Identifying coral to genus level is difficult and requires a long period of training to acquire a high degree of accuracy. To address this issue, Frontier Cambodia Marine has created key morphological categories to attain increased identification accuracy during substrate surveys (Table 5). It is suggested that the three dimensional physical structure of the reef has a bigger role in influencing reef fish assemblage than the overall health of the coral (Yahya et al., 2011). Friedlander and Parish (1998) proposed that a more complex habitat may provide additional refuge from predators and currents, as well as increasing the availability of resources. The key structural coral categories created could be used as a proxy for coral reef complexity e.g. branching coral has a more complex three-dimensional structure than massive coral Methodology Coral and substrate surveys occurred along transects set up for surveying reef fish abundance (Table 3). After completing the reef fish survey, five locations were randomly selected along each 20 m transect. A 50 by 50 cm quadrat was placed at each of these locations and used to assess the percentage cover for each substrate category (Table 5). Any damage to coral in the form of bleaching, anchor damage or disease was also recorded. If any dead coral was observed, it was recorded even if it was not possible to ascertain causation. After completing each of the five sample quadrats, end depth was measured and end time recorded. 15

16 Table 5. Categories used to assess coral cover and substrate type on reefs surrounding Koh Rong Samloem Category Massive coral Sub-massive coral Branching coral Meandering coral Plates of coral Corals with daytime polyps Corals forming columns Solitary corals Sea fan Sea rod Sea plume Sponge Coralline branching algae Coralline encrusting algae Macroalgae - Fleshy Macroalgae - Filamentous Sand Bare rock Rubble Litter Other information to be recorded Percentage cover of each coral category present will be recorded. If there is bleaching, damage to coral or diseases (type) it will also be recorded. Type of litter will be recorded 16

17 Mean ± SE percentage cover of dominant substrates found on House Reef transects Mean ± SE percentage cover of dominant substrates found on House Reef transects CMM 142 Science Report Results Substrate Type Figure 8. Mean ± SE percentage cover of dominant substrates found at Transect 1-6 on House Reef during phase Massive Submassive Algae Sand Rock Substrate Type Figure 9. Mean ± SE percentage cover of dominant substrates found at Transect 1-6 on House Reef during phase

18 Consistent with observations made during phase 141, House Reef is dominated by filamentous algae; massive and submassive coral, rock and sand (Fig. 8). Other coral structures, such as branching, meandering, plate, daytime exposed polyps and solitary are present in lower frequencies and make up a small proportion of the substrata. There were some differences identified between the data sets collected from House Reef s transects during phase 141 versus phase 142 (Fig. 8 and Fig. 9). Table 6. Mean ± SE percentage cover massive corals and algae on House Reef transects Massive coral Algae Transect 1 (141) (± 4.75 SE) 37.7 (± 5.53 SE) Transect 2 (141) 41.4 (± 6.87 SE) 37.7 (± 6.23 SE) Transect 3 (141) (± 5.90 SE) 24 (±4.75 SE) Transect 4 (141) (± 6.83 SE) 27.1 (± 5.5 SE) Transect 1 (142) 31.9 (± 1.02 SE); 30.3 (± 1.07 SE). Transect 2 (142) (± 5.82 SE) 33.8 (± 4.76 SE) Transect 3 (142) (± 5.5 SE); 20.9 (±3.43 SE). Transect 4 (142) (± 5.45 SE) 21.8 (± 3.86 SE). During phase 141, transects 1-4 all showed increasing proportions of massive coral and similarly decreasing proportions of filamentous algae and submassive coral (Fig. 9). Data collected during phase 142 follows similar patterns in respect to trends between categories, although individual transects show variation between the two sets (Table 6). The two most notable differences between the data sets are the decline in the percentage of rock observed along transects and the presence of dead coral, previously undocumented on House Reef. Rock was only found on transects 2 and 3 during phase 142, with mean percent substrate cover recorded at 1.33 (±1.33 SE) and 1.8 (±1.8 SE) respectively. Dead coral was observed in low frequency on all House Reef transects except transect 2 where it was not observed at all. Transect 1; 2.67 (±1.0 SE), transect 3; 3.3 (±2.1 SE), transect 4; 1.13 (±0.94 SE), transect 5; 3.67 (±2.46 SE) and transect 6; 4.89 (±2.52 SE) Discussion While reef fish abundance and diversity are of primary importance to the stakeholders of Koh Rong Samloem, it is important to assess coral cover and composition for a complete look at the status of a reef. Coral cover and complexity are both factors that influence fish abundance. Studies demonstrating that 18

19 there is a positive relationship between live coral cover and the total number of reef fish individuals (Bell and Galzin, 1984; Feary et al., 2009) were supported by findings during phase 141; the site with higher mean percentages of live coral cover also had higher mean total fish abundance. Reef complexity, or the physical structure of the coral, also increases the biodiversity potential of the area through creating ecological niches and is important to reef fish (Bell and Galzin, 1984; Friedlander and Parish, 1998; Yahya et al., 2011), which was also in agreement with last phase s findings. Differences in substrate composition observed between the two phases include a decrease in the percentage of rock along the transects and the occurrence of dead coral, which was previously not observed during surveys. Again, these divergences are likely a result of sampling effect as this project is still in the early phases of data collection. However, substrate surveys for the next phase will closely monitor the frequency and proportion of dead coral on House Reef to verify whether or not it is increasing, as it could indicate a serious problem for the condition of the reef. Cheal et al. (2002) state that fish diversity gives a poor estimate of a reef s resilience to a disturbance event, highlighting the need to assess reef substrate composition in areas prone to disturbances. Given the current, rapid increases in tourist development in the M pai Bay area, it is essential that future monitoring covers both reef fish assemblage and coral habitat Invertebrate surveys Introduction Invertebrate species are an important component of coral reef ecosystems and provide a wide variety of ecosystem services. They are important grazers, filter feeders and predators within the reef community and perform key services to reef systems in the form of nutrient recycling, water quality regulation and herbivory (Przeslawski et al., 2008). In addition to fish and benthic substrate surveys, invertebrate diversity and abundance surveys were conducted to sample the coral reef systems surrounding Koh Rong Samloem as a whole. Invertebrates play an important role in reef health and due to their vulnerability to climate change (Hutchings et al., 2007), it is important to obtain baseline data of invertebrates inhabiting the reefs off Koh Rong Samloem Methodology Transects set up for surveying reef fish abundance and benthic composition were also used to assess the abundance of benthic invertebrates (Table 7). After completing the reef fish survey, but before measuring the end depth of the transect, the abundance and diversity of invertebrates present were recorded. Five stations were randomly selected along each 20 m transect. At each station a 50 by 50 cm quadrat was placed. At each of these five stations, any benthic invertebrates present within the quadrat area were recorded according to the categories shown in Table 7. 19

20 Table 7. Categories of invertebrates surveyed during the fish and benthic composition surveys Groups of invertebrates recorded in surveys Worms Bivalves Feather duster worm Christmas tree worm Flatworm Crustaceans True crab Shrimp spp. Anemone shrimp Mantis shrimp Hermit crabs Gastropods Topshell Cowrie Conch Drupella spp. Cone shell Tridacna spp. Sea pen Cephalopods Octopus Echinoderms Chocolate drop starfish Cushion star Feather star Crown of thorns starfish Brittlestar Collector urchin Pencil urchin Diadema spp. Holothuria spp. Synaptid spp. Murex shell Nudibranch spp. Phyllidia spp. Jorunna funebris Glossodoris spp. Phyllidiella spp. 20

21 Mean ± SE abundance of invertebrates found at House Reef CMM 142 Science Report Results Consistent with the last phase, the most common invertebrates on House Reef were clams; 2.93 (± 0.45 SE), Diadema; 0.94 (±0.23 SE), feather duster worms; 0.70 (± 0.2 SE) and Drupella 0.45 (± 0.2 SE). Benthic species outside of these were rarely or never sighted Invertebrate type Figure 10. Mean ± SE abundance of invertebrates found at House Reef. Clams were the most common type of invertebrate found on House Reef. As seen below in Fig. 11, transect 5 has the greatest number of clams (3.4 ± 0.52 SE) while transect 3 had significantly fewer clams than any of the other transects (1.83± 0.41 SE). The distribution of the other types of common invertebrates was mostly consistent across House Reef transects, with the exceptions of significantly more Drupella observed on transect 2 (1.43± 0.05 SE) and significantly less Diadema observed on transect 3 (0.13± 0.24 SE). There was no significant difference between the number of feather duster worms on any given transect. 21

22 Mean ± SE abundance of the most common invertebrates found at House Reef CMM 142 Science Report Transect clam feather duster worm drupella diadema Figure 11. Mean ± SE abundance of common invertebrates found at House Reef Discussion Clams have been the most commonly observed type of invertebrate on House Reef during both phase 141 and 142. Clams and other bivalves are filter feeders and play an important part in recycling nutrients (Przeslawski et al., 2008). This is crucial for reef systems as they form in oligotrophic waters and need to be very efficient in their use of nutrients (Jackson and Buss, 1975). Feather duster worms, another of the commonly observed invertebrates, are also filter feeders and participate in this process as well. Personal observations by staff and volunteers indicate that giant clams, sea pens and other bivalves have been harvested and empty shells discarded onto the reef. Growth of the tourism industry as well as an expanding local population may lead to increased bivalve fishing, which would reduce the amount of filter feeding and exacerbate the nutrient load in the water. Coupled with increased sewage outflow from wastewater sources, this could shift the current conditions towards a system that has increased algal cover and thus, decreased coral cover. This would likely reduce species diversity and may negatively impact fish abundance, as well as increase the reef s vulnerability to disturbances such as cyclones or sea surface temperature rises. 4. Proposed work programme for next phase The proposed work programme for next phase includes a continuation of the current species surveys: fish, coral and invertebrate surveys on House Reef and Sunset Rock Reef, with an emphasis on collecting data from Sunset if tidal levels permit. Other potential survey sites will be investigated and integrated into the program as appropriate. During the next phase a proposal to conduct socio-economic survey work will be developed and, if approved, work will commence on developing a survey questionnaire to be put to the local population. 22

23 5. References Bell, J. D., & Galzin R. (1984). Influence of live coral cover on coral-reef fish communities. Marine Ecology Progress Series. 15: Cambodian Ministry of Tourism, (2013). Available at statistic_reports/tourism_statistics_annual_report_2013.pdf. Date visited June Carpenter, K. E., & V. G. Springer. (2005). The center of the center of marine shore fish biodiversity: The Philippine islands. Environmental Biology of Fishes 72: Cheal, A. J., Coleman, G., Delean, S., Miller, I., Osbourne, K., & Sweatman H., (2002). Responses of coral and fish assemblages to a severe but short-lived tropical cyclone on the Great Barrier Reef, Australia. Coral Reefs 21: Chou, L. M., Tuan, V. S., Philreefs, Yeemin, T., Cabanban, A., Suharsono, & Kessna, I., (2002). Status of Southeast Asia Coral Reefs. Status of Coral Reefs of the World. Davenport, J., & Davenport, J.L. (2006). The impact of tourism and personal leisure transport on coastal environment: A review. Estuarine, Coastal and Shelf Science, 67, De Lopez, T. T. (2003). Economics and stakeholders of Ream National Park, Cambodia. Ecological Economics. 46: Feary, D. A., McCormick, M. I., & Jones G. P., (2009). Growth of reef fishes in response to live coral cover. Journal of Experimental Marine Biology and Ecology 373: Friedlander, A. M., & Parrish, J. D., (1998). Habitat characteristics affecting fish assemblages on a Hawaiian coral reef. Journal of Experimental Marine Biology and Ecology 224: 1-30 Hawkins, J. P., & Roberts, C. M. (1994). The growth of coastal tourism in the Red Sea: present and future effects on coral reefs. Ambio. 23: Hutchings, P., Ahyong, S., Byrne, M., Przeslawski, R. and G. Wörheide, G. (2007). Chapter 11: Vulnerability of benthic invertebrates of the Great Barrier Reef to climate change. In Climate change and the Great Barrier Reef: a vulnerability assessment. Great Barrier Reef Marine Park Authority Home Page at: Accessed June 2014 Jackson, J. B. C., & Buss, L. (1975). Allelopathy and spatial competition among coral reef invertebrates. Proceedings of the Natural Academy of Science USA. 72: Jennings, S., Brierley, A. S., & Walker, J. W. (1994). The inshore fish assemblages of the Galapagos Archipelago. Biological Conservation. 70: Jones, G. P., McCormick, M. I., Srinivasan M., & Eagle, J. V. (2004). Coral decline threatens fish biodiversity in marine reserves. Proceedings of the National Academy of Sciences 101: Killeen, T. J. (2012). The Cardamom Conundrum: Reconciling development and conservation in the Kingdom of Cambodia. NUS Press, National University of Singapore. 23

24 Lewis, A. R. (1998). Effects of experimental coral disturbance on the population dynamics of fishes on large patch reefs. Journal of Experimental Marine Biology and Ecology 230: Przeslawski, R., Ahyong, S., Byrne, M., Worheide, G., & Hutchings P. (2008). Beyond corals and fish: the effects of climate change on noncoral benthic invertebrates of tropical reefs. Global Change Biology. 14: Reef Check, (2007). Monitoring Protocol. Available at: Date visited January 2014 Touch, S. T. (1995). Present status of fisheries in Cambodia and the development action plans. The Department of Fisheries. Phnom Penh. Wilkinson, C., & Souter D. (2008). Status of Caribbean coral reefs, bleaching and hurricanes in Global Coral Reef Monitoring Network and Reef and Rainforest Research Centre, Townsville, Australia. Williams, D. M. (1982). Patterns in the distribution of fish communities across the Central Great Barrier Reef. Coral Reefs. 1: Wilson, S. K. Graham N. A. J., Pratchett M. S., Jones G. P., & Polunin N. V. C. (2006). Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient? Global Change Biology 12: Yahya, S. A. S., Gullstrom, M., Ohman, M. C., Jiddawi, N. S., Andersson, M. H., Mgaya, Y. D., & Lindahl, U. (2011). Coral bleaching and habitat effects on colonistation of reef fish assemblages: An experimental study. Estuarine, Coastal and Shelf Science 94: Zhao, J., Neil, D. T., Feng, Y., Yu, K., & Pandolfi, J. M. (2009). High-precision U-series dating of very young cyclone-transported coral reef blocks from Heron and Wistari reefs, southern Great Barrier Reef, Australia. Quaternary International 195: