managing the impacts of scuba divers on thailand's coral reefs

19
Managing the Impacts of SCUBA Divers on Thailand’s Coral Reefs Suchai Worachananant School of Natural and Rural Systems Management, University of Queensland St Lucia Campus, Brisbane, Australia R.W. (Bill) Carter Faculty of Science, Health and Education University of the Sunshine Coast, Queensland, Australia Marc Hockings School of Natural and Rural Systems Management, University of Queensland Gatton Campus, Australia Pasinee Reopanichkul Centre for Water Studies, School of Engineering, University of Queensland St Lucia Campus, Australia While dive tourism enjoys continued growth worldwide, concern exists that it is con- tributing to the degradation of coral communities, biologically and aesthetically. This study examined the effect of SCUBA diver contacts with coral and other substrates. Ninety-three percent of divers made contact with substrata during a 10-minute obser- vation period with an average of 97 contacts per hour of diving. Two-thirds of the divers caused some coral damage by breaking fragments from fragile coral forms with an average of 19 breakages per hour of diving. Fin damage was the major type of damage. Underwater photographers caused less damage per contact than non-photographers; as did male divers, compared with females. Diver-induced damage decreases with increasing number of logged dives and attendance at pre-dive briefings. Park man- agers can help reduce impact by identifying and directing use to sites that are resistant to damage, matching diver competence and site preferences, and alerting operators to dive conditions. Minimising impact requires dive operators to be proactive in promot- ing minimal impact diving behaviour. This includes selecting sites that match diver expectations and experience, and providing pre-dive briefings in the context of diver activities and physical capacity, and site susceptibility to impact and current strength. doi: 10.2167/jost771.0 Keywords: coral reefs, education, environmental impact, management response, SCUBA divers, Surin Marine National Park 0966-9582/08/06 645-19 $20.00/0 C 2008 Taylor & Francis JOURNAL OF SUSTAINABLE TOURISM Vol. 16, No. 6, 2008 645 Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Upload: uq

Post on 01-May-2023

0 views

Category:

Documents


0 download

TRANSCRIPT

Managing the Impacts of SCUBA Divers onThailand’s Coral Reefs

Suchai WorachananantSchool of Natural and Rural Systems Management, University ofQueensland St Lucia Campus, Brisbane, Australia

R.W. (Bill) CarterFaculty of Science, Health and Education University of the SunshineCoast, Queensland, Australia

Marc HockingsSchool of Natural and Rural Systems Management, University ofQueensland Gatton Campus, Australia

Pasinee ReopanichkulCentre for Water Studies, School of Engineering, University ofQueensland St Lucia Campus, Australia

While dive tourism enjoys continued growth worldwide, concern exists that it is con-tributing to the degradation of coral communities, biologically and aesthetically. Thisstudy examined the effect of SCUBA diver contacts with coral and other substrates.Ninety-three percent of divers made contact with substrata during a 10-minute obser-vation period with an average of 97 contacts per hour of diving. Two-thirds of the diverscaused some coral damage by breaking fragments from fragile coral forms with anaverage of 19 breakages per hour of diving. Fin damage was the major type of damage.Underwater photographers caused less damage per contact than non-photographers;as did male divers, compared with females. Diver-induced damage decreases withincreasing number of logged dives and attendance at pre-dive briefings. Park man-agers can help reduce impact by identifying and directing use to sites that are resistantto damage, matching diver competence and site preferences, and alerting operators todive conditions. Minimising impact requires dive operators to be proactive in promot-ing minimal impact diving behaviour. This includes selecting sites that match diverexpectations and experience, and providing pre-dive briefings in the context of diveractivities and physical capacity, and site susceptibility to impact and current strength.

doi: 10.2167/jost771.0

Keywords: coral reefs, education, environmental impact, managementresponse, SCUBA divers, Surin Marine National Park

0966-9582/08/06 645-19 $20.00/0 C© 2008 Taylor & FrancisJOURNAL OF SUSTAINABLE TOURISM Vol. 16, No. 6, 2008

645

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

646 Journal of Sustainable Tourism

IntroductionHealthy coral reefs make many contributions to coastal communities. They

help lessen the erosive impacts of waves in the coastal zone, support the liveli-hoods of fishing communities and provide the opportunity for many coastalcommunities to enhance their incomes from the rapid expansion of globaltourism (Sudara et al., 1991; Dixon, 1993; Wilkinson et al., 1994; Seenprachawong,2003). Thailand’s maritime environments are a major attraction for internationalvisitors: more than 80% of overseas tourists come to Thailand to visit Thailand’sseas (Seenprachawong, 2001; Thamrongnawasawat & Worachananant, 2004). Atpresent, there are more than 20 million visitors travelling to the seas of Thailandeach year (Worachananant & Thamrongnawasawat, 1999; Piewsawat, 2002; TAT,2006). This is due, at least in part, to the diversity of organisms in Thai reefs(Sudara, 2002a).

While coral reefs are one of the most popular resources for tourist use (Hall,2001; Sudara, 2002b), there is increasing concern for the impact tourist activitymay be having on reefs. The impacts of recreation activities have received atten-tion from many marine researchers (Woodland & Hooper, 1977; Kay & Liddle,1986, 1989; Liddle & Kay, 1987; Sudara et al., 1991; Hawkins & Roberts, 1992;Mohamaed et al., 1994; Rouphael & Inglis, 1997; Al-Jufaili et al., 1999; Rouphael& Inglis, 2001; Tratalos & Austin, 2001). The activities that were identified tobring about changes include reef walking, snorkelling, development of facilitiesand pollution of waters by powerboats. Some studies have raised the concernthat SCUBA diving may also constitute a significant threat (Rouphael & Inglis,2002; Davenport & Davenport, 2006).

Breakage of corals by SCUBA divers has been documented worldwide, in-cluding reports from Egypt (Hawkins & Roberts, 1992), the USA (Talge, 1992),Australia (Rouphael & Inglis, 1997) and the Caribbean (Tratalos & Austin, 2001).Divers damage corals through direct physical contact with their hands, body,equipment and fins (Rouphael & Inglis, 1995, 2001; Barker & Roberts, 2004).Some damage may be from diving-associated activities, such as anchoring,rather than solely from diver-induced damage (Jameson et al., 1999). The dam-age caused by individual divers is often considered minor (Walters & Samways,2001; Zakai & Chadwick-Furman, 2002), although there is some evidence thatthe cumulative effects of these disturbances can cause significant localised de-cline of coral cover (Hawkins et al., 1999). A study in the Red Sea also found thatthe percentage of hard coral cover decreased by 43% and algal cover increasedover four-fold (Jameson et al., 2007). While most divers contact corals duringtheir dive, it has been reported that the majority of divers appear to have littleimpact, with only a few divers causing substantial damage (see Harriott et al.,1997; Rouphael & Inglis, 1997; Walters & Samways, 2001; Zakai & Chadwick-Furman, 2002).

There is evidence that the characteristics of individual divers, such as level ofdive education and briefing, diving experience, gender, and camera possession,affect the number of contacts with coral and the amount of damage (Rouphael &Inglis, 1995, 2001). Medio et al. (1997) report that a single environmental aware-ness briefing resulted in a reduced rate of diver contacts with reef substances,with voluntary contacts mainly directed at the non-living substrate. Roberts and

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 647

Harriot (1995) found that divers with more advanced levels of training tend tohave fewer impacts. Moreover, male divers appear to cause significantly moredamage because men tend to be more adventurous, less likely to follow in-structions and more likely to take risks than women (Rouphael & Inglis, 1997).Rouphael and Inglis (2001) found that use of a camera had no influence on therate or amount of damage caused by divers. Studies have found that the level ofdamage caused to coral communities does vary with the experience of SCUBAdivers, and the proportion of vulnerable types of organisms (e.g. branchingcorals), but damage is not clearly related to reef topography (Rouphael & Inglis,1997; Zakai & Chadwick-Furman, 2002). There are numbers of studies world-wide that have examined diver experience in relation to dive site characteristics(Chanwichai, 1994; Dearden et al., 2006; Kubas et al., 2006; Musa et al., 2006;Worachananant et al., 2006). The main contributors to diving satisfaction includethe quality of the underwater nature landscape, the comfort and ease of accessto dive sites, distance and the cost of accommodation and food. Some studiesfound a relationship between dominant coral types and diver satisfaction. Siteswith a high proportion of fragile coral types received higher satisfaction fromdivers (Worachananant et al., 2006; Chanwichai, 1994). However, sites domi-nated by fragile corals might not be preferred dive sites by managers becauseof their vulnerability to physical impact. There is a need to balance the needs ofboth divers and site managers. If the diving tourism industry is to sustain itself,there is a need to limit diver-induced stress on the coral reef system. Because ofthe great variation in the amount of physical damage within and among div-ing sites (Harriott et al., 1997; Jameson et al., 1999; Worachananant et al., 2006),management and restoration efforts should be conducted on a site-by-site basis.

The aim of this study was to examine the impact on coral caused by SCUBAdivers to Surin Marine National Park in Thailand, and to determine if the topog-raphy of coral reef dive sites influences the type and amount of damage causedby divers. The study involved observation of the behaviour of SCUBA divers,with impacts examined in relation to diver characteristics, and the character ofreefs used by divers, including the percentage cover and the dominant life formof the corals. The objective was to examine the extent to which issues affectingthe impact of divers on reefs in other studies apply to Surin and to consider themanagement options that could help limit impacts while still providing appro-priate diving experiences for visitors to the park. Specific null hypotheses thatwere tested are:

(1) pre-dive briefing does not affect the rate of contact between divers andsubstrates and the extent of damage resulting from these contacts;

(2) use of an underwater camera does not influence the extent of damage causedby divers;

(3) there is no difference between male and female divers in their rates of contactwith, and damage to, the substrates; and

(4) there is no difference in the contact and damage rates of divers based ontheir diving experience.

These elements of diver behaviour provide insights for the development of op-tions for managing divers’ use of reef resources in ways that minimise impacts.Knowledge of the biological factors that affect the sensitivity of different sites to

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

648 Journal of Sustainable Tourism

SCUBA diver impacts can be combined with knowledge of diver characteristicsthat make them more likely to cause coral damage. They can be combined inorder to allow managers to match diver characteristics with suitable locationsso as to minimise diver impacts while delivering satisfying and sustainableexperiences.

The Study Area: Surin Marine National ParkThe Thailand Ministry of Natural Resources and Environment administers

the 135 km2 Surin Marine National Park (hereafter, Surin). Seventy-six percentof the area (102 km2) is marine, including 8 km2 of reef, and the balance (33 km2)is terrestrial. Surin consists of five granitic islands (North Surin, South Surin,Torinla, Pachumba and Stork Islands) and two exposed pinnacles (Figure 1).

Figure 1 Islands and reefs of Surin Marine National Park

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 649

More than 68 species of corals are found in Surin and the condition of thereefs is considered good (DMSCI, 1997; Dearden et al., 2000; Worachananantet al., 2004; Worachananant et al., 2005). The pollution-free clear water supportshealthy coral growth, especially in the sheltered bays on the eastern sides ofthe islands such as Turtle Bay and Mae Yai Bay. Torinla Island, the area wherethe reef is in best condition, has more than 90 percent living coral coverage.Corals generally extend from low water to a depth of 30 metres. In shelteredareas, seabeds deeper than 30 metres are sandy. Encrusting corals are found inexposed locations, especially on the western side of the park where strong wavesbatter the shore during the monsoon season. Here, the seabed below 10 metresis mainly sandy with a few rocks.

In common with most reefs in Thailand, reef condition at Surin has declined inthe past decade (Worachananant et al., 2007). El Nino effects, particularly in 1998,caused deterioration of reefs in some areas, especially Mae Yai Bay through coralbleaching. However, the park is remote from the mainland (around 60 km) andall the islands are covered with healthy forest, so sedimentation and pollutionare not major issues. Most researchers suggest that the major threat is fromhuman-related activities (Thamrongnawasawat et al., 2000; Sittithaweepat, 2001;Saisaeng, 2002; Worachananant et al., 2007).

Survey MethodsThe study involved observation of 108 SCUBA divers, selected by convenience

from dive parties visiting five different dive sites at Surin (Mai Ngam Bay,Pakkhad Bay, Turtle Bay, Torinla Island and Suthep Bay [Figure 1]) betweenNovember and December 2004, which is the peak season for diving at Surin.This sample represents just over one quarter of the 402 SCUBA divers that parkrecords showed as visiting Surin during November and December 2004. All ofthe divers in the sample are Thai (nearly 95% of park visitors are Thai nationals),and most are female (an ad hoc survey of eight dive operators working inThailand indicated that females normally represent about two-thirds of theclients of dive trip operations). Surveys of visitors to Mu Koh Surin NationalPark in December 2004 and March 2005 (Worachananant, 2007) also reporteda positive female gender balance amongst visitors to the park (62% and 60%,respectively). The reasons behind the predominance of female clients of diveoperations in Thailand are not known, but it appears that the diver sample inthis study is not abnormal in this regard. After December 2004, the number ofSCUBA divers dropped sharply following the 2004 Boxing Day Tsunami (TAT,2005a, 2005b), with only about 50 divers in January 2005 compared to nearly1,000 in January 2004 (Worachananant et al., 2006).

Selection aimed for a representative sample of the demographics of divegroups, including age, gender, physical characteristics and level of diving expe-rience. The study was conducted on three diving vessels from two companieswho agreed to support the study. Daily selection of dive sites by the local divestaff was based on a number of criteria, including weather and sea conditions,popularity of the site (i.e. whether it was requested by visiting divers), and therelative experience of visiting divers. At most sites, diving was restricted to amaximum depth of 15 metres. As part of their pre-dive briefing, the dive staff

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

650 Journal of Sustainable Tourism

would describe the location of ‘points of interest’ within the dive area, suchas manta-ray or whale shark, explain the plan of the dive, including directionand dive time, and remind divers to be careful not to touch or break livingorganisms, especially corals. This pre-dive briefing session was part of the touroperator’s standard procedure.

Each subject was observed underwater for 10 minutes. The dives usuallylasted about 60 minutes and divers were observed during only one of the10-minute periods of the dive. Information was recorded underwater on thegender of each diver, the site where they were observed, the time of observa-tion during the dive (i.e. which of the 10-minute periods) and whether theycarried a camera. For each subject, the level of training and experience (numberof dives completed since gaining qualifications) were obtained from the divestaff onboard. Observers were introduced to the diving group as normal divers;they entered the water with the dive party, and remained a distinct distancebehind their subjects (usually 5–10 metres depending on visibility). To avoidinfluencing the subjects’ behaviour, divers were not informed of the activitiesof the observers until all dives were completed. Data from subjects who wereobviously aware of the presence of the observers were not used. Each diver whomade multiple dives in the trip was recorded only once; their other dives wereexcluded from the sample.

Three observers participated in this study. To minimise bias between ob-servers, comprehensive training in data collection methods and criteria wasconducted at the beginning of each field trip.

The number of times that each diver made direct physical contact with thesubstratum, broken or damaged corals during the dive was recorded. Contactand damage were classified according to whether they were made by the diver’shands, fins, knees, gauges or other equipment and the type of substratum in-volved. Benthic substrata were categorised as branching (including tabulate),massive (including sub-massive), foliose and encrusting corals, and other sub-strates. For completeness, the disturbance to ‘other substrates’ was recordedwhen sessile organisms other than hard corals (e.g. soft corals or seafans) weredamaged, sand was disturbed or rocks were dislodged.

The mean number of diver interactions resulting in coral damage in each10-minute period of diving was compared by one-way ANOVA. Independentt-tests were used to assess the difference between the mean number of dam-aging contacts and various attributes (camera possession, gender and pre-divebriefing status). Chi-square tests were used to determine the characteristics ofdivers who damaged corals. Correlation between level of divers’ experienceand number of divers who made contact with or damaged corals was analysedusing Spearman’s rank correlation coefficient.

During the period of the diver observations, living coral coverage datawere assessed using line intercept transects based on methods described inWorachananant et al. (2007). All transects were located in the edge-slope zone(between 4 and 10 metres depth): the standard zone used for this type of monitor-ing in Thailand (Phongsuwan & Chansang, 1992; DMSCI, 1997; DMCR, 2005).Percentage cover was calculated using the length of the transect interceptedby each of the coral forms/substrates. Differences between coral type cover atdifferent dive sites were tested by analysis of variance. Coral life forms were

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 651

divided into branching, massive, foliose, encrusting, sub-massive, tabulate anddead. Non-coral areas were recorded as ‘other substrates’ and included sand,rock and other benthic organisms. To interpret the results, percentage coralcover was subsequently arranged into three categories: ‘tolerant’ coral types(massive, sub-massive and encrusting), ‘fragile’ types (branching, tabulate andfoliose) and ‘dead and others’ (dead coral and other substrates).

Results

Diver characteristics and their impact on coral reefsObservation subjects were mainly female (77%). Diving experience varied

widely; the number of logged dives ranged from 4 to 560 (median = 27 dives).Subjects were placed into four groups based on experience (Table 1). Around70% of divers had logged fewer than 50 dives before arriving at Surin.

One hundred and one divers (94%) made some contact with the substratesduring the 10-minute observation period. Divers averaged 16.2 ± 1.9 (mean ±SE) contacts per 10 minutes (approximately 97 times per hour’s dive). Of thosedivers who made contact with the substrate, 82 divers (81%) made contact withsome form of coral with an average of 5.5 ± 0.7 contacts per 10 minutes (33 timesper dive).

Seventy-one divers (66%) damaged coral at least once during the 10-minuteobservation period. On average, contact with the substratum that resulted incoral damage occurred around 3.1 ± 0.4 per 10 minutes (19 breakages per dive).Visible damage caused by the divers normally consisted of the breaking offragments of branching corals. The damage to massive and encrusting coralswas not clearly seen, but the generation of mucus by affected coral was evidentand recorded as damage (Liddle & Kay, 1987). Kicks by divers’ fins were themajor cause of coral damage (Table 2).

Of the many types of substrates, branching coral and other substrates suchas rock, sand and dead coral received the highest level of damage (193 and 168records, respectively). However, the proportion of contact and damage on eachsubstrate varied. Foliose corals and branching corals were the most vulnerable tophysical impact. All foliose corals and 75 percent of branching corals contactedby divers were damaged (Figure 2).

Thirty-nine divers (52%) who attended a pre-dive briefing (‘attending divers’)damaged corals. A significantly higher proportion (32 divers or 97%) who didnot attend the pre-dive briefing (‘non-attending divers’) caused damage (Table 3;

Table 1 Number of divers’ logged dives

Groups Frequency Percent

Beginners (1–25 dives) 44 41

Novices (26–50 dives) 29 27

Enthusiasts (51–100 dives) 8 7

Experts (more than 100 dives) 27 25

Total 108 100

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

652 Journal of Sustainable Tourism

Table 2 Number of times divers contacted coral reef substrates

Types Coral Other Totalsubstrate

Branching Massive Foliose Encrusting

Fin 151 112 7 31 176 477

Hand 51 95 2 20 191 359

Gauges 34 22 25 13 30 124ContactKnee 0 0 0 0 111 111

Other 21 9 0 0 52 82

Total 257 238 34 64 560 1153

Fin 108 15 7 21 101 252

Hand 35 30 2 11 38 116

Gauges 34 8 25 8 20 95DamageKnee 0 0 0 0 1 1

Other 16 3 0 0 8 27

Total 193 56 34 40 168 491

Figure 2 Number of times divers contacted and damaged substrates

χ2(1,106) = 20.58, p < 0.01). In addition, the average number of corals damaged by

non-attending divers was significantly greater than damage caused by attendingdivers (Figure 3; t(1,106) = −3.32, p < 0.01). A Levene’s test of homogeneity ofvariance also suggests that the variance in damage rate of non-attending diverswas higher than for attending divers (F(1,106) = 5.61, p < 0.05). That is, if diversattend a pre-dive briefing, they are likely to have fewer contacts with coral andcause less physical damage.

A significantly smaller percentage of non-photographers caused damage com-pared to divers with cameras (Table 3: χ2

(1,106 ) = 7.03, p < 0.05). The average num-ber of corals damaged by underwater photographers was not significantly dif-ferent to that caused by non-photographers (t(1,106) = −0.74, p > 0.05) (Figure 3).

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 653

Figure 3 Mean number of contacts and damage incidents (±SE) over 10 minutes anddiver characteristics

Table 3 Impact on corals and diver characteristics

Contact Damage Coralwith coral to coral damage

afterNumber % Number % contact (%)

Attended pre-dive briefing

No (n = 33) 32 97 32 97 100

Yes (n = 75) 50 67 39 52 78

Camerapossession

No (n = 51) 30 59 27 53 90

Yes (n = 57) 52 91 44 77 85

Gender Female (n = 83) 67 81 62 75 93

Male (n = 25) 15 60 9 36 60

Level ofexperience

1–25 dives (n = 44) 39 89 35 80 90

26–50 dives (n = 29) 19 66 15 52 79

51–100 dives (n = 8) 8 100 7 88 88

100 dives up (n = 27) 16 59 14 52 88

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

654 Journal of Sustainable Tourism

A Levene’s test of homogeneity of variance suggests that the variance in damagerate of non-photographers is greater than for photographers (F(1,106) = 5.04, p <

0.05). That is, the number of non-photographers who damaged corals was fewerthan photographers, but the damage per contact rate was higher.

A significantly greater percentage of female divers (75%, n = 62) than maledivers (36%, n = 9) damaged corals (χ2

(1,106) = 12.78, p < 0.05). The averagedamage rate per contact by female divers was also significantly greater thanthat caused by male divers (Figure 3; t(1,106) = 5.28, p < 0.05). A Levene’s testof homogeneity of variance also suggests that the variance in damage rate offemale divers was greater than male divers (F(1,106) = 25.68, p < 0.05). That is,females are more likely to make contact with corals and cause more damagethan males.

There was a correlation between the level of experience (number of loggeddives) and the number of times divers came into contact with and damagedcorals (r = −0.34, n = 108, p < 0.05 and r = −0.31, n = 108, p < 0.05, respectively).This indicates that the number of times divers damage corals decreases withincreasing level of diving experience (Figure 3).

To test whether the difference in damage caused to corals was simplyexperience-related rather than gender-related, analysis revealed that there wasno difference in the experience range by gender of divers who damaged corals(ANOVA, F(3,104) = 4.29, p > 0.05). That is, whether expert or novice, femaledivers are more likely to cause damage to corals than males.

Seventy-six percent of all contacts that resulted in damage to corals were‘uncontrolled contacts’ (interactions where the diver did not purposely touchthe substrata; see Davis & Tisdell, 1995) caused by a fin kick (47%), gauge (23%)and other parts (6%). Beginner and novice divers are more likely to make contactwith corals than more experienced divers (ANOVA, F(3,104) = 7.07, p < 0.01). Allother contacts (24%) were caused by hand (i.e. divers consciously touched orgrabbed corals). Again, novice divers are more likely to do this than experienceddivers (ANOVA, F(3,104) = 4.80, p < 0.01).

Biological differences between diving sitesMean cover of corals from five selected dive sites differed greatly (Table 4).

While the reef at Torinla Island is dominated by branching corals, the reef atTurtle Bay is dominated by tolerant coral types.

Table 4 Coral cover at the five dive sites

Study sites Coral cover (%)

Tolerant Fragile Dead and other

Mean SE Mean SE Mean SE

Torinla Island 9.17 2.12 76.18 2.73 14.66 1.87

Pakkhad Bay 22.95 1.99 60.32 2.03 16.73 2.01

Suthep Bay 33.44 1.51 42.57 2.15 23.98 1.71

Mai Ngam Bay 37.34 2.37 26.23 2.42 36.43 1.66

Turtle Bay 47.42 2.47 23.26 1.81 29.32 2.20

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 655

Figure 4 Substrate contacts per diver (mean ± SE) over a 10-minute interval at the fivedive sites

Mean cover of all coral forms, except tabulate coral, varied significantly be-tween sites. Mean cover of tolerant forms (combining massive, sub-massive andencrusting corals) ranged from 9.2 ± 2.1% (mean ± SE) at Torinla Island to47.4 ± 2.5% at Turtle Bay. Cover of tolerant coral differed greatly among sites(ANOVA, F(4,125) = 40.79, p < 0.01). A Tukey HSD test indicates that percentcover means of the tolerant corals of Torinla Island, Pakkhad Bay and TurtleBay differ significantly, while the mean of the tolerant corals cover at Mai NgamBay did not differ from Suthep Bay. Cover of fragile corals ranged from 23.3 ±1.8% at Turtle Bay to 76.2 ± 2.7% at Torinla Island. The mean cover of fragilecorals differed significantly between sites (F(4,125) = 91.81, p < 0.01). A TukeyHSD test indicates that the means of the percent cover of fragile corals at TorinlaIsland, Pakkhad Bay and Suthep Bay differed significantly, while mean fragilecoral cover at Mai Ngam Bay did not differ from Turtle Bay.

While the strength of the currents at offshore islands such as Surin is related totidal change/exchange of water (Phongsuwan & Chansang, 1994), the differentgeographical characteristics of each diving site influences the strength of current.Torinla Island, a channel site, has very strong currents, while Pakkhad Bay andTurtle Bay, which are exposed bays, have medium currents during ebb tide. MaiNgam Bay and Suthep Bay are almost enclosed bays and experience little tidalcurrent. Torinla Island has a much higher rate of contact than the other sites,while differences between the other sites were less marked (Figure 4).

Difference in diver behaviour among dive sitesSubjects frequently came into contact with coral at all five dive sites. The

mean number of contacts per 10 minutes ranged between 7.3 and 37.0 times perdiver (Figure 4). There was a significant difference between the mean number ofcontacts at different dive sites (ANOVA, F(4,103) = 13.38, p < 0.01), with TorinlaIsland, the site with the strongest current, accounting for a much higher numberof contacts.

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

656 Journal of Sustainable Tourism

Figure 5 Contacts (%) resulting in damage at the five dive sites

Table 5 Damage caused by divers over a 10-minute period at the five dive sites

Study sites N Mean SE Minimum Maximum

Torinla Island 24 8.50 1.25 0 21

Pakkhad Bay 24 2.33 0.65 0 12

Suthep Bay 24 1.21 0.47 0 9

Mai Ngam Bay 24 1.25 0.28 0 6

Turtle Bay 12 1.00 0.39 0 4

Total 108 3.06 0.44 0 21

Twenty-two divers (92%) observed at Torinla Island made at least one contact(in a 10-minute interval) with corals. Of this, 21 divers (88%) caused damage.Turtle Bay was the least affected site, with only 10 divers (42%) causing damageto corals (Figure 5). The total number of contacts that resulted in damage rangedfrom 4 at Turtle Bay to 21 at Torinla Island. The mean number of contacts thatresulted in damage was larger at Torinla Island (8.5 ± 1.25) than at the other foursites (Table 5).

DiscussionIn this study, SCUBA divers often made contact with corals, and on more

than half of these occasions coral was damaged. The amount of damage and thefrequency of its occurrence were relatively high (around 19 times per dive). Theresults suggest that divers practice a low level of environmental care. However,divers may simply be unaware that their actions can injure coral.

A short pre-dive presentation on environmental protecting dive behaviourwas shown to reduce the damage caused by divers. Divers who attended apre-dive briefing were likely to have fewer contacts with corals and cause less

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 657

physical damage than those who did not attend briefings (i.e. null hypothesis 1is rejected). During these briefings, dive staff reminded divers to be careful notto touch and break living organisms, especially corals. Photographers touchedcorals more often than non-photographers and thus damaged more per dive,but damage per contact was less. That is, photographers tended to be gentlerwhen they made contact with coral, but this level of care is insufficient to reducethe net effect of the contacts (null hypothesis 2 is rejected). Female divers weresignificantly more likely to damage corals when compared with males (nullhypothesis 3 is rejected). This may be due to differences in physical capacityand ability to handle the heavy SCUBA equipment and currents, rather thanany difference in care or concern. The number of times novice divers cameinto contact with corals, whether uncontrolled or conscious, was higher thanfor more experienced divers and diver-induced damage to corals decreasedwith increasing number of logged dives (null hypothesis 4 is rejected). Table 6compares our findings with the literature on diver-induced impacts on coral.

Site-Related EffectsThe amount of damage caused by divers was also related to the percent cover

of fragile corals, which were the most vulnerable to physical impact. This meansthat sites with a high cover percent of fragile coral are more vulnerable to in-creasing pressure of divers than places with a high proportion of tolerant corals.The site with the strongest current also showed the highest level of damage tocorals. While places with a high percent cover of fragile coral are more vulnera-ble, factors such as diver strength, experience and activity undertaken stronglyinfluence the degree of damage. Female divers, with less physical strength thanmales, are more likely to make contact with the coral substrate because of theirlower physical capacity to handle currents and bulky diving gear. Some activ-ities, such as underwater photography, can increase diver-induced damage tocoral.

Conceptually from this study, the site factors that influence SCUBA diverpropensity to cause damage to coral can be placed on two axes: coral type andcurrent (Figure 6). Current in this case represents the physical characteristics ofthe water column and possibly includes other water characteristics that affectdiver comfort and stability, including turbidity. Low body strength, the absenceof pre-dive briefing and photography are factors that can decrease the rangeof sites made available to divers if concern for impact is given high priority. Ifminimising diver attrition of coral is desired, then beginner divers should beconfined to sites dominated by resistant coral species and low current strength(the northwest sector of Figure 6). As divers have increasing experience, sitesdominated by fragile coral types and stronger currents can be made availablewith less risk of damage to the corals.

Reducing the damage to dive sites, especially in areas with strong current anda high proportion of fragile corals (the southwest sector of Figure 6), involvesconsiderations relating to body strength or fitness, the nature of any specialactivity undertaken while diving (e.g. photography), and the level of awarenessdivers have of their potential to damage. While, from a management perspective,little can be done to regulate diver strength, body mass (gender correlated

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

658 Journal of Sustainable Tourism

Table 6 Comparison of this study with findings in the literature

Issue Literature This study Possible explanation

Attendanceat pre-divebriefingreducingdiver-induceddamage tocorals

Medio et al. (1997)and Davis andTisdell (1995) foundthat a pre-divebriefing led to areduction in impact,but Barker andRobert (2004) foundno effect.

Divers attendinga pre-divebriefing havefewer contactswith coral andcause lessphysicaldamage.

Most literature supports ourfinding that pre-dive briefingcan reduce diver-induceddamage to corals.A one-sentence inclusion in aregular dive briefing askingdivers to avoid touching thereef might not be sufficient toraise the conservationawareness of divers (seeBarker & Robert, 2004).

Camerapossessionaffects theextent ofdiver-induceddamage.

Literature reportsmixed results, withBarker and Robert(2004) finding thatphotographers hadsignificantly highercontact rates, whileRouphael and Inglis(2001) found noeffect.

Photographers(divers withcamera) touchedcoral more oftenthan non-photographersand thusdamaged moreper dive, butdamage percontact was less.

Underwater cameras arebecoming a more affordableaccessory for many divers,both expert and novice. Oneof the authors (SW), anexperienced divinginstructor, suggests thatnovice divers who havelimited buoyancy control aremore likely to contact coralswhen using a camera.

Genderdifferencesin diver-induceddamage tocoral.

Rouphael and Inglis(1997) found thatmale divers causesignificantly moredamage, but femaledivers were morelikely to hold or totouch benthicsubstrata (Rouphael& Inglis, 2001).

Females aremore likely tomake contactwith coral andcause damagethan males.

Our finding is likely to be theresult of differences inphysical capacity and abilityto handle the heavy SCUBAequipment and currentsrather than any difference incare or concern.

Experience-andtraining-relateddifferencesin diver-induceddamage tocoral.

Literature reportsmixed results.Roberts andHarriott (1995)found that diverswith moreadvanced levels oftraining have fewerimpacts, althoughHarriott et al. (1997)and Rouphael andInglis (2001) foundno significantdifferences in thetotal numbers ofcontacts or impactsmade by divers ofdifferent levels ofexperience.

Number of timesdivers damagecorals decreaseswith increasinglevel of divingexperience.Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 659

Figure 6 Factors influencing SCUBA diver impacts to coral

diver characteristics) and fitness, there is potential to facilitate best practiceunderwater photography techniques and increasing awareness of the potentialfor divers to do damage. These require an education programme delivered eitherby park staff or by dive-boat operators.

The framework presented in Figure 6 can also be used as an input to zoningplans. While other factors will influence the zoning of SCUBA diving use zones,sites with attributes on the eastern side of the figure are likely to be acceptableto stakeholders for comfort and safety reasons and therefore make managementeasier.

Conclusion: Improving the Sustainability of SCUBA DivingAt an extreme, diver impact could be minimised by focusing all SCUBA div-

ing in resistant sites that are current free. Throughout the world, and especiallytourism-dependent Thailand, such action would be unacceptable to users, op-erators and management and would conflict with park management objectives.However, within the framework presented (Figure 6), there is considerable roomfor flexibility. The largest proposed constraint is water characteristics (e.g. cur-rent strength): a constraint that will be acceptable to divers because safety andcomfort are desired by recreational SCUBA divers. Equally desired by divers,especially by experienced divers, are sites with a high proportion of the morefragile coral types (e.g. branching and plate coral types). For park managementthen, the task is to identify and facilitate SCUBA diving at sites that match diverexpectations for safety and quality experiences, but within the experience of the

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

660 Journal of Sustainable Tourism

Table 7 SCUBA diving stakeholder actions to minimise contact impact

Park management actions Dive-boat operator, dive master anddive-club actions

Define dive sites with marked mooringbuoys indicating recommendedexperience levels (largely determined bycoral substrate type).

Promote minimal impact divingtechniques in all promotional material.

Develop partnerships with dive mastersand dive-boat operators to communicatebest practice diving technique thatincludes minimum impact diving.

Include information on potential forsubstrate-contact in all pre-dive briefings.

Provide information on diver impactsgenerally and especially to dive mastersand dive-boat operators.

Give greater emphasis to this if the diversare beginners and novices, the dive site isrich in fragile corals, dive conditions arenot benign, or divers intend to undertakeactivities that demand interaction withthe substrate (e.g. photography).

Provide minimal impact divinginstruction to dive masters anddive-boat operators.

Inform divers (dive boat operators) ofdive conditions daily, and recommenddive sites for divers of different levels ofexperience.

diver to dive with minimum impact (the eastern sectors of Figure 6). Practically,such an approach can only be achieved with the co-operation of divers, diveoperators and dive masters. Therefore, park managing agencies must take thelead in building partnerships with key stakeholders to implement such a policy.For the partnership to succeed, all stakeholders must make a commitment toa shared vision. Actions that will demonstrate commitment are presented inTable 7.

While implementing environmental/minimal impact awareness programmesfor divers is recommended in many reef areas, these have largely been conductedonly by park managing agencies (Medio et al., 1997). As dive-based tourism isan important revenue earner for many resource users, and if the sites are tomaintain their aesthetic appeal and biological characteristics, then it is in theinterests of dive operators to ensure their clients dive safely and with minimalimpact. While protected area managers could take draconian zoning action tominimise impacts by focusing use away from sensitive areas, this would have theeffect of reducing the attractiveness of Surin (as the example), and clients wouldprobably move elsewhere. It is suggested that collaborative action is a moresensible, and indeed a more effective and responsible approach. Park managersneed to identify suitable diving sites with stakeholders and provide data suchas presented here to inform selection of diving sites. Then, a coordinated divereducation programme that informs divers of their responsibilities is needed tocontinually deliver a minimal impact diving message. Diving federations and

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 661

associations, such as PADI, NAUI or CMAS, and tour operators, as well asindividual dive schools and instructors can probably take a lead in this.

CorrespondenceAny correspondence should be directed to Suchai Worachananant, De-

partment of Marine Science, Kasetsart University, 50 Phaholyothin Road,Chatuchuk, Bangkok, Thailand 10900 ([email protected]).

ReferencesAl-Jufaili, S., Al-Jabri, M., Al-Baluchi, A., Baldwin, R.M., Wilson, S.C., West, F. and

Matthews, A.D. (1999) Human impacts on coral reefs in the Sultanate of Oman. Estu-arine, Coastal and Shelf Science 49 (Suppl. A), 65–74.

Barker, N.H.L. and Roberts, C.M. (2004) Scuba diver behaviour and the management ofdiving impacts on coral reefs. Biological Conservation 120, 481–489.

Chanwichai, D. (1994) Visitors profile: The tourism attitude toward Surin Marine Na-tional Park, Thailand. MS thesis, James Cook University, Queensland.

Davenport, J. and Davenport, J.L. (2006) The impact of tourism and personal leisuretransport on coastal environments: A review. Estuarine, Coastal and Shelf Science 67(1–2), 280–292.

Davis, D. and Tisdell, C. (1995) Recreational SCUBA-diving and carrying capacity inmarine protected areas. Ocean and Coastal Management 26, 19–40.

Dearden, P., Bennett, M. and Rollins, R. (2006) Implications for coral reef conservation ofdiver specialization. Environmental Conservation 33, 353–363.

Dearden, P., Theberge, M. and Bennett, M. (2000) Monitoring and marine park man-agement at Koh Surin and Mu Koh Similan, Thailand. Paper presented at ManagingProtected Areas in a Changing World: Proceedings of the Fourth International Con-ference on Science and Management of Protected Areas, 14–19 May 2000, Wolfville,N.S.

Dixon, J.A. (1993) Economic benefits of marine protected areas. Oceanus 36, 35–40.DMCR (2005) Assessment report on effects of tsunami to coastal resources in Andaman

sea, Thailand (in Thai). Department of Marine and Coastal Resources (DMCR), Min-istry of Natural Resources and Environment with collaboration of 9 Alliance of MarineScience Institutes of Thailand.

DMSCI (1997) Mu Koh Surin Marine National Park Survey Report (in Thai). Unpublisheddocument, Department of Marine Science (DMSCI), Faculty of Fisheries, KasetsartUniversity, Bangkok, Thailand.

Hall, C.M. (2001) Trends in ocean and coastal tourism: The end of the last frontier? Oceanand Coastal Management 44 (9–10), 601–618.

Harriott, V., Davis, D. and Banks, S.A. (1997) Recreational diving and its impact in marineprotected areas in Eastern Australia. Ambio 26, 173–179.

Hawkins, J.P. and Roberts, C.M. (1992) Effects of recreational SCUBA diving on fore-reefslope communities of coral reefs. Biological Conservation 62, 171–178.

Hawkins, J.P., Roberts, C.M., Van’t Hof, T., de Meyer, K., Tratalos, J. and Aldam, C.(1999) Effects of recreational scuba diving on Caribbean coral and fish communities.Conservation Biology 13, 888–897.

Jameson, S.C., Ammar, M.S.A., Saadalla, E., Mostafa, H.M. and Riegl, B. (1999) A coraldamage index and its application to diving sites in the Egyptian Red Sea. Coral Reefs18, 333–339.

Jameson, S.C., Ammar, M.S.A., Saadalla, E., Mostafa, H.M. and Riegl, B. (2007) A quan-titative ecological assessment of diving sites in the Egyptian Red Sea during a periodof severe anchor damage: A baseline for restoration and sustainable tourism manage-ment. Journal of Sustainable Tourism 15, 309–323.

Kay, A.M. and Liddle, M.J. (1986) The impact of reefwalking at Hardy Reef. GBRMPA reportstatus.

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

662 Journal of Sustainable Tourism

Kay, A.M. and Liddle, M.J. (1989) Impact of human trampling in different zones of acoral-reef flat. Environmental Management 13, 509–520.

Kubas, A., Altas, D. and Sapci, B. (2006) Analysis of the motivation factors that affect thedivers’ diving place preferences by the ordinal logistic regression. International Journalof Agriculture and Biology 8.

Liddle, M.J. and Kay, A.M. (1987) Resistance, survival and recovery of trampled coralson the Great Barrier Reef. Biological Conservation 42, 1–18.

Medio, D., Ormond, R.F.G. and Pearson, M. (1997) Effect of briefings on rates of damageto corals by scuba divers. Biological Conservation 79, 91–95.

Mohamaed, M.I.H., Rahman, R.A. and Abdullah, M.P. (1994) Impact of developmentof Pulau Layang Layang coral reefs. In C.R. Wilkinson, S. Sudara and L.M. Chou(eds) Third ASEAN-Australia Symposium on Living Coastal Resources (Vol. 2, pp. 35–40).Townsville, Australia: Australian Institute of Marine Science.

Musa, G., Kadir, S.L.S.A. and Lee, L. (2006) Layang Layang: An empirical study onSCUBA divers’ satisfaction. Tourism in Marine Environments 2, 89–102.

Phongsuwan, N. and Chansang, H. (1992) Assessment of coral communities in the An-daman Sea (Thailand). Paper presented at the 7th International Coral Reef Symposium,Guam, Micronesia, 22–27 June 1992.

Phongsuwan, N. and Chansang, H. (1994) Long term monitoring of coral reefs in theAndaman Sea, Thailand. In: C.R. Wilkinson, S. Sudara and L.M. Chou (eds) ThirdASEAN-Australia Symposium on Living Coastal Resources (Vol. 2, pp. 5–12). Townsville,Australia: Australian Institute of Marine Science, Thailand.

Piewsawat, W. (2002) Analysis of recreation activities and effectiveness of nature inter-pretation program at Mu Koh Surin national park, Phang-Nga province. MS thesis,Kasetsart University, Thailand.

Rouphael, A.B. and Inglis, G.J. (1995) The effects of qualified recreational SCUBA diverson coral reefs. Technical report no. 4, CRC Reef Research Centre Ltd., Townsville,Australia.

Rouphael, A.B. and Inglis, G.J. (1997) Impacts of recreational SCUBA diving at sites withdifferent reef topographies. Biological Conservation 82, 329–336.

Rouphael, A.B. and Inglis, G.J. (2001) Take only photographs and leave only footprints?:An experimental study of the impacts of underwater photographers on coral reef divesites. Biological Conservation 100, 281–287.

Rouphael, A.B. and Inglis, G.J. (2002) Increased spatial and temporal variability incoral damage caused by recreational scuba diving. Ecological Applications 12, 427–440.

Saisaeng, A. (2002) Taxonomic study on fish family Pomacentridae in Thai Coral Reef.MS thesis, Kasetsart University, Thailand.

Seenprachawong, U. (2001) Capturing coral reef benefit values: Financing marine envi-ronment conservation at Phi Phi Islands, Thailand. Paper presented at Asian WetlandSymposium 2001, Penang, Malaysia.

Seenprachawong, U. (2003) Economic valuation of coral reefs at Phi Phi Islands, Thailand.International Journal of Global Environmental Issues 3, 104–114.

Sittithaweepat, N. (2001) Study of species richness and distribution of Nudibranchs inThai Coral Reef. MS thesis, Kasetsart University, Thailand.

Sudara, S. (2002a) Conservation and management of MPA in Thailand. Paper presented atThe Fourth Conference on the Protected Areas of East Asia. IUCN World Commissionon Protected Areas in East Asia Conference ‘Benefits Beyond Boundaries in East Asia’,Taipei, Taiwan, 18–23 March, 2002.

Sudara, S. (2002b) Current status and prospect of protected areas in Thailand. Paperpresented at The Fourth Conference on the Protected Areas of East Asia. IUCN WorldCommission on Protected Areas in East Asia Conference “Benefits Beyond Boundariesin East Asia”, Taipei, Taiwan, 18–23 March, 2002.

Sudara, S., Thamrongnawasawat, T., Nateekanjanalarp, S. and Kuanman, P. (1991) Coralreef management plan for conservation and tourism development in Ang Tong, Samuiand Pha Ngan Islands in the Gulf of Thailand. Paper presented at The RegionalSymposium on Living Resource in Coastal Areas, University of Philippines, Manila.

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009

Impacts of SCUBA Divers on Thailand’s Coral Reefs 663

Talge, H. (1992) Impact of recreational divers on scleractinian corals at Looe Key, Florida.Paper presented at the Seventh International Coral Reef Symposium, Guam, 22–26June 1992.

TAT (2005a) New strategy for the recovery of tourism after tsunami. On WWW at www.world-tourism.org/tsunami/news/58.pdf. Accessed 22.11.05.

TAT (2005b) Tsunami news. On WWW at http://www.tatnews.org. Accessed 5.10.05.TAT (2006) Tourism Statistic. On WWW at http://www2.tat.or.th/stat/web/static index.

php. Accessed 5.5.06.Thamrongnawasawat, T. and Worachananant, S. (2004) Mu Koh Surin Marine Na-

tional Park. On WWW at http://www.talaythai.com/issue/surin park/index.php3.Accessed 29.11.04.

Thamrongnawasawat, T., Worachananant, S., Saisaeng, A., Sittithaweepat, N.,Limviriyakul, P. and Patimanukasem, O. (2000) Survey report in Mu Koh Surin MarineNational Park Area presented to UNESCO project. Unpublished paper.

Tratalos, J.A. and Austin, T.J. (2001) Impacts of recreational SCUBA diving on coralcommunities of the Caribbean island of Grand Cayman. Biological Conservation 102,67–75.

Walters, R.D.M. and Samways, M.J. (2001) Sustainable dive ecotourism on a South Africancoral reef. Biodiversity and Conservation 10 (12), 2167–2179.

Wilkinson, C.R., Sudara, S. and Soekarno (1994) Socio-economic values and impactson ASEAN coral reefs. In C.R. Wilkinson, S. Sudara and C.L. Ming (eds) ThirdASEAN-Australia Symposium on Living Coastal Resources (Vol. 1, pp. 23–31). Townsville,Australia: Australian Institute of Marine Science.

Woodland, D.J. and Hooper, J.N.A. (1977) The effect of human trampling on coral reefs.Biological Conservation 11, 1–4.

Worachananant, S. (2007) Management approaches in marine protected areas: A casestudy of Surin Marine National Park, Thailand. PhD thesis, The University of Queens-land, Australia.

Worachananant, S., Carter, B. and Hockings, M. (2006) Managing tourism in Surin MarineNational Park, Thailand. Paper presented at the East Asian Seas Congress 2006, HaikouCity, Hainan Province, China, 12–16 December 2006.

Worachananant, S., Carter, B., Hockings, M. and Reopanichkul, P. (2006) Reef recoveryand management in Surin Marine National Park. Paper presented at Post-DisasterAssessment and Monitoring of Coastal Ecosystems, Biological and Culture Diversityin The Indian Ocean and Asianwaters, Phuket, Thailand, 20–23 February 2006.

Worachananant, S., Carter, B., Hockings, M., Reopanichkul, P. and Thamrongnawasawat,T. (2004) Tourism management in Surin Marine National Park, Thailand. Paper pre-sented at Coastal Zone Asia Pacific Conference ‘Improving the Quality of Life inCoastal Areas’, Hilton Hotel, Brisbane, Australia, 5–9 September 2004.

Worachananant, S., Carter, B., Hockings, M., Reopanichkul, P. and Thamrongnawasawat,T. (2005) Management response to the tsunami, Surin Marine National Park, Thailand.Paper presented at the 1st international Marine Protected Areas Congress, WaterfrontCampus, Deakin University, Geelong, Australia.

Worachananant, S., Carter, R.W. and Hocking, M. (2007) Impacts of the 2004 tsunami onSurin Marine National Park, Thailand. Coastal Management 35, 399–412.

Worachananant, S. and Thamrongnawasawat, T. (1999) Pilot study of Gorgonians dis-tribution in Thai Sea. Paper presented at Kasetsart University Annual Conference,Kasetsart University, Main Campus, Bangkok, Thailand.

Zakai, D. and Chadwick-Furman, N.E. (2002) Impacts of intensive recreational diving onreef corals at Eilat, northern Red Sea. Biological Conservation 105, 179–187.

Downloaded By: [University of the Sunshine Coast] At: 08:29 23 November 2009