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Parrotfish bioerosion on Egyptian Red Sea reefs

Article  in  Journal of Experimental Marine Biology and Ecology · April 2009

DOI: 10.1016/j.jembe.2009.01.019

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 ARTICLE IN PRESS
JEMBE-48808; No of Pages 7
Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx

Contents lists available at ScienceDirect

Journal of Experimental Marine Biology and Ecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / j e m b e

1 Parrotfish bioerosion on Egyptian Red Sea reefs

2 Magdy A. Alwany a,b,c,⁎, Ellen Thaler b, Michael Stachowitsch c
3 a
Department of Marine Science, Faculty of Sciences, Suez Canal University, Ismailia, Egypt
4 b
Institute of Zoology and Limnology, University of Innsbruck, Innsbruck, Austria
5 c

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Faculty of Life Sciences, Department of Marine Biology, University of Vienna, Althanstrasse 14, A-1090, Vienna, Austria
^
6

a r t i c l e i n f o a b s t r a c t
7
8 Article history:

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Parrotfishes (family Scaridae) are important agents in marine bioerosion. Here, the feeding ecology of seven
9 Received 30 March 2008 species of parrotfishes was studied on Egyptian Red Sea reefs. The most abundant species on both the reef flat
10 Received in revised form 9 August 2008 and slope was Chlorurus sordidus. In contrast, C. gibbus had the lowest abundance on the reef flat, and
11 Accepted 29 January 2009
Cetoscarus bicolor was the least abundant species on the reef slope. Scarus niger exhibited the highest
12 Available online xxxx
feeding rate (98.9 bites 5 min− 1), followed by C. sordidus (76.5 bites 5 min− 1), whereas the rates for C. bicolor
13 ^ ^
16 and C. gibbus were low (29.4 and 31.9 bites 5 min− 1, respectively). The daily feeding patterns of all seven
Keywords: ^
17
18
19
20
21
Bioerosion
Corallivory
Coral reefs
Egypt
Feeding
^
DP
species showed agreement in that activity was relatively constant over the day, with highest values in the
early afternoon (1400 h) and a steady decrease until 1800 h. C. sordidus was more similar to S. niger and
^
S. ghobban in showing somewhat higher activities in the morning (0800 h) followed by a slight decrease until

3
^
noon. The average bite volumes of C. gibbus and C. bicolor were high (0.114 and 0.110 cm3, respectively),
whereas S. niger had the lowest average value (0.002 cm ). Based on their feeding intensity, C. gibbus,
22 Parrotfishes S. ghobban and C. bicolor have high bioerosion rates on the Egyptian Red Sea reefs. Overall, S. ghobban is the
23 Red Sea
most important bioeroder because it is more abundant than the other two species. All parrotfish species fed
TE
37 on dead coral and hard substrates which are rich in algae, but C. gibbus, C. bicolor and S. ghobban also fed on
38 live coral on both reef zones; C. sordidus avoided live coral. The fresh scars on live coral were bigger than on
39 dead coral because the three large parrotfish (C. gibbus, C. bicolor and S. ghobban) fed mainly on live corals.
40
41 © 2009 Published by Elsevier B.V.
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42

45 1. Introduction Quantitative estimates of parrotfish bioerosion, however, are 63

largely restricted to the Caribbean (Gygi, 1969, 1975; Ogden, 1977; 64
46 Bioerosion is a major structuring force operating on coral reefs
RR

Scoffin et al., 1980; Bruggemann et al., 1994a,b, 1996), and Australian 65

47 (Hutchings, 1986) and may be categorized as either external (grazing, ^
reefs (Bellwood, 1995a,b, 1996). Previous studies that aimed to 66
48 e.g. by fishes) or internal (boring, e.g. by sponges). It is the removal of quantify parrotfish bioerosion on different reefs have usually 67
49 material from the reef matrix by biological processes. The first note on employed indirect methods (except: Bruggemann et al., 1994a,b, 68
50 bioerosion by parrotfishes was by Darwin (1842, 1845). Because ^
1996 in the Caribbean Sea and Bellwood, 1995a,b, 1996 in the Great 69
51 of their abundance, parrotfishes (Scaridae) are important agents of Barrier Reef), measuring total ingestion and estimating the proportion 70
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52 marine bioerosion (Bellwood, 1995a,b). As a dominant component of reworked sediments. In the Red Sea, gut content analyses have been 71
53 of the herbivore assemblage on coral reefs, many parrotfish species successfully carried out in the past (Al-Hussaini, 1945, 1947; Gohar 72
54 have specialized in scraping algae off limestone; other species can ^
and Latif, 1959, 1961). The present study uses direct methods to 73
55 excavate reef substrates and harvest substrate-bound algae (Horn, quantify scarid bioerosion based on grazing scars on reefs of the 74
^
56 1989). Choat (1991) reported that the piscine bioerosion may be an Northern Egyptian coast of the Red Sea. This study is also the first 75
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57 important agent of structural change on reefs (e.g. sediment recycling attempt to calculate the bioerosion rate of parrotfishes on Red Sea 76
58 by taking it up directly from reef aprons). This importance may even reefs by examining daily feeding activity and abundance. The role of 77
59 exceed that of sea urchins on some well-developed coral reefs in the seven parrotfish species was evaluated and the bioerosion rate on 78
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60 central Red Sea (Hassan, 1997). Sea urchins may be responsible for 80% different spatial scales (reef zones, sites and regions) was determined. 79
61 of the total bioerosion of reefs in some regions and fishes have the
62 high percentage in other regions (Scoffin et al., 1980). 2. Materials and methods 80

⁎ Corresponding author. Department of Marine Science, Faculty of Sciences, Suez

Canal University, Ismailia, Egypt. Tel.: +20 10 4499116.
E-mail address: magdy.elalwany@yahoo.com (M.A. Alwany).
Observations were made while snorkeling and/or SCUBA diving 81
^ ^
over the reef flat (RF) and reef slope (RS) at water depths from 1–8 m. 82
^
Notes were taken on PVC slates and underwater paper and data were 83

0022-0981/$ – see front matter © 2009 Published by Elsevier B.V.

doi:10.1016/j.jembe.2009.01.019

Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
jembe.2009.01.019
 ARTICLE IN PRESS
2 M.A. Alwany et al. / Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx

84 collected while following individual fish at a distance of 3–5 m. This 2.2. Studied scarid species 122
^ ^
85 approach minimized interference with natural behaviour while still
86 permitting the collection of accurate data. The field work was carried The total scarid fauna in the Northern Egyptian coast of the Red Sea 123
87 out during daylight hours (0800–1900 h); where feeding activities consists of approximately 22 species (Goren and Dor, 1994). The seven 124
^ ^
88 were measured every 2 h interval (0800, 1000, 1200, 1400, 1600 and species selected here were chosen based on preliminary observations 125
^ ^
89 1800 h), and abundances were counted between 1100 and 1300 h. of the potential role in bioerosion according to size and feeding 126
^ ^
90 Data were gathered during several field trips conducted during one activity. The selected species are: Cetoscarus bicolor (Bicolour parrot- 127
91 season (early summer of 2000) on the Egyptian Red Sea reefs. fish), Chlorurus gibbus (Steepheaded parrotfish), C. sordidus (Bullet- 128
head parrotfish), Scarus ferrugineus (Rusty parrotfish), S. frenatus 129
92 2.1. Study area (Bridled parrotfish), S. ghobban (Bluebarred parrotfish), and S. niger 130
(Swarthy parrotfish). The studied species therefore include the full 131
93 All field observations were undertaken on the reefs in three range of adult scarid sizes, i.e. from the relatively small S. niger 132
94 regions of the Northern Egyptian coast of the Red Sea. Each region (SL 12.6–22.5 cm) to large-sized species such as C. gibbus (SL 41.5 cm) 133
^^ ^
95 includes two sites, a and b, which were chosen randomly for the (sizes after Khalaf and Disi, 1997). They were identified according to 134
^
96 behavioural observations (Fig. 1). Moreover, the reef flat and the Randall (1983) and based on personal communication with him 135
97 reef slope were distinguished for certain parameters (e.g. abun- (2002). 136
98 dance, number of scars on live and dead corals). First region

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99 includes sites 1a and b; site 1a (27° 50′ N– 34° 19′ E): is located at 2.3. Abundance of scarid bioeroders 137
^^
100 Ras Um Elseid reefs in Sharm El-Sheikh city. The reef flat here is
^
101 relatively poor and extends over a width of 50–70 m. The reef edge Parrotfish abundance was quantified by means of visual censuses, 138
^
102 and slope exhibit well-developed coral colonies. Site 1b (27° 43′ N– as the most common, non-destructive method to quantify fish 139
^ ^ ^
103 34° 14′ E): is located at Yolanda reefs in the southern tip of Sinai abundances (Sale, 1980). Species were counted visually along 100 m 140

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104 Peninsula, inside the Ras Mohamed National Park (20 km south of long, 5 m wide and 1 m high transects (100 × 5 × 1 = 500 m3) laid on the 141
105 site 1a). The reef flat is very rich in corals and represents a model for reef flat and reef slope parallel to the shoreline. 142
106 well-developed reefs. Second region includes sites 2a and b; site 2a
^
107 (28° 36′ N– 34° 30′ E): is a famed diving site located north of Dahab 2.4. Feeding rates and mean daily bites 143
^^
108 city in the Gulf of Aqaba. The branched corals on the reef edge
109
110
111
112
consist mainly of Acropora. The reef flat here is between 20 and
40 m wide. Site 2b (28° 32′ N– 34° 29′ E): is located off El-Masbat
^^
drops steeply to 20 m depth, then gently after that. The poorly
^
Police Station in Dahab city (6 km south of site 2a). The reef slope
^ DP Foraging behaviour was quantified by following individual fish for
a specific time and tallying the number of bites taken from each
substratum item. Feeding rate (= number of bites per time), as the
index of feeding activity, was determined for all seven species. The
144
145
146
147
113 developed reef flat is narrow in the northern part (10 m) and wide feeding study included a total of 214 parrotfish. Each species was 148
114 in the southern part (50 m). Third region includes sites 3a and b; studied at each of the 6 sites. At each site, the activity of 4 individuals 149
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115 site 3a (27° 13′ N– 33° 56′ E) consists of a relatively small reef patch of each species was observed: for each individual fish, bites on various 150
^^
116 (about 250 m2). The reef flat has scattered branched coral colonies substrata were counted for three consecutive periods of 5 min, with 1– 151
^
117 (mostly Acropora spp.). The slope around the patch is gentle (depth 2 min intervals (in some cases, less than 4 individuals of C. gibbus were 152
^ ^
118 ranging between 5 and 11 m). Site 3b (27° 14′ N– 33° 58′ E) lies recorded). Observations were discontinued if the individual was lost 153
^^
119 1200 m north of site 3a. This reef extends far into the water (900 m) during the 5 min observation period or if the individual showed any 154
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120 and contains very well- developed coral colonies throughout, signs of disturbance. To estimate daily feeding patterns, the feeding 155
^
121 including the reef edge and slope. rates were recorded during a series of snorkel surveys from 0800 to 156
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Fig. 1. Map of Egypt and the northern part of the Red Sea coast showing the study sites.

Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
jembe.2009.01.019
 ARTICLE IN PRESS
M.A. Alwany et al. / Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx 3

157 1900 h on several days [morning (0800–1000 h), noon (around Table 2 t2:1
^ ^ ^ Feeding rates (bites 5 min− 1), bite volumes (cm3) and bioerosion rates (cm3 h− 1 and kg h− 1)
158 1200 h) and afternoon (1400–1600 h)].
^ ^ ^ of seven parrotfish species in the Northern Red Sea
t2:2
159 2.5. Bioerosion rates Species Feeding rates Bite volumes Bioerosion rates t2:3
bites/5 min cm3 3 −1
cm h kg h −1
t2:4
160 Bite areas and volumes were estimated according to Bellwood Cetoscarus bicolor 29.4 ± 4.2 0.110 ± 0.066 40.8 ± 27.7 0.0571 t2:5
161 (1995a) by following individual fish until the exact location of a bite Chlorurus gibbus 31.9 ± 9.9 0.114 ± 0.062 47.0 ± 32.1 0.0672 t2:6
162 could be clearly seen. The observer swam to the bite site and C. sordidus 76.5 ± 13.2 0.008 ± 0.003 7.0 ± 2.8 0.0098 t2:7
Scarus ferrugineus 59.4 ± 11.9 0.009 ± 0.008 6.9 ± 8.3 0.0097 t2:8
163 measured the size of the bite using vernier callipers. The maximum S. frenatus 53.6 ± 5.4 0.011 ± 0.005 7.1 ± 3.9 0.0100 t2:9
164 width, length and depth of the feeding scar were measured to the S. ghobban 54.6 ± 8.7 0.063 ± 0.025 41.2 ± 16.7 0.0577 t2:10
165 nearest 0.1 mm. A total of 30 bites were examined for each species at S. niger 98.9 ± 7.7 0.002 ± 0.001 2.5 ± 0.9 0.0035 t2:11
166 each site.
167 After a slight modification of Bellwood (1995a), bioerosion rates
168 were calculated as follows: on the reef slope at most sites. ANOVA showed that no significant 200
differences between both reef zones (Table 1). 201
−1
Mean bioerosion rate ðcm3 h Þ = estimated mean bite rate


ðbites per hourÞ×mean bite volume cm3 : 3.2. Feeding activity 202

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170
171 The daily bite rate was calculated by multiplying the mean bite rate Scarus niger had the highest feeding rate among all species (98.9 bites 203
172 per hour by 12 (which represented the day length during the early 5 min− 1), followed by C. sordidus (76.5 bites 5 min− 1), whereas C. bicolor 204
^ ^ ^ ^
173 summer study period). As day length during winter may drop below and C. gibbus had the lowest values (29.4 and 31.9 bites 5 min− 1, 205
^
174 10 h and increase to 14 h during summer, this represents a reasonable respectively in Table 2). Three species had nearly similar feeding rates: 206

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^ ^
175 average. Annual bioerosion rates were calculated by multiplying daily S. frenatus (53.6), S. ghobban (54.6) and S. ferrugineus (59.4). The studied 207
176 rates by 365 (Bellwood, 1995a; Bruggemann et al., 1996). Bioerosion species can therefore be divided into three categories based on feeding 208
^
177 volumes were converted into weight equivalents based on the skeletal rates. The first includes S. niger and C. sordidus, which have high feeding 209
−3
178 density of 1.4 g cm given by Hughes (1987; cited in Bak, 1990) for rates. The second includes three species (S. ferrugineus, S. ghobban and 210
^
179 Porites species — the species on which bite volumes were calculated in S. frenatus) with moderate rates. The last one includes C. bicolor and 211
^
180
181
182
183
the present study. Bites were considered to be shallow, rounded,
elongate depressions and individual bite volumes calculated based on
geometric formulae in http://mathworld.wolfram.com/CircularSeg-
ment.html.
DP C. gibbus, which have low rates. The ANOVAs showed that the feeding
rates did not differ significantly between the sites (Table 3).
The daily feeding patterns on the Northern Red Sea reefs are shown
in Fig. 2. The activities of all species were relatively constant over the
day. The patterns of C. sordidus, S. niger and S. ghobban show certain
212
213
214
215
216
184 2.6. Parrotfish scars on corals similarities [higher in the morning (from 0800 to 1000 h), slight 217
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decrease at noon (around 1200 h), increase in the afternoon (from 218
^
185 Food selection was quantified for each fish individual. The number 1400 to 1600 h) and slight drop before sunset]. Although there was no 219
^
186 of bites on live and dead corals was counted in 6 replicates for 5 min, clear trend of daily feeding for the other parrotfish species, the values 220
^
187 and the average per 5 min was then calculated. A total of 427 also tended to be higher in the early afternoon and then steadily drop 221
^
188 individuals of seven parrotfish species were observed on both the reef until 1800 h. 222
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189 flat and slope.

3.3. Parrotfish bioerosion 223
190 3. Results
The average bioerosion rates expressed as volumes and weights 224
191 3.1. Abundance of bioeroding parrotfish per time are summarized in Table 2. Excluding S. niger (due to its very 225
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low bioerosion rate), the other species were divided into two distinct 226
192 The most abundant species was Chlorurus sordidus on both reef groups. The first is characterized by high bioerosion rates and includes 227
193 zones (reef flat and slope, Table 1). In contrast, C. gibbus had the lowest C. gibbus, S. ghobban and C. bicolor (N40.8 cm3 h− 1). The second is 228
^ ^ ^ ^
194 values on the reef flat, while C. bicolor had the lowest values on the characterized by low rates (b7.1 cm3 h− 1) and includes S. frenatus, 229
^ ^
195 reef slope. Certain species were almost equally abundant in both reef C. sordidus and S. ferrugineus. According to Bellwood and Choat (1990), 230
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196 zones at all sites, e.g. C. bicolor, S. niger and S. frenatus. C. gibbus and we can name the first group as excavators and second group as 231
^
197 S. ghobban displayed distinctive zonation patterns on reef habitats: scrapers. The species of the second group exhibited very similar rates. 232
198 C. gibbus preferred the reef slope and S. ghobban the reef flat. Combining this information with the abundance data, S. ghobban and 233
199 Generally, however, the number of parrotfish individuals was higher C. sordidus are probably the main contributors to overall bioerosion 234
rates on the Northern Red Sea fringing reefs based on their high 235
abundance. The ANOVAs showed that the bioerosion rates did not 236
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differ significantly between the sites (Table 3). 237

t1:1 Table 1
Average abundance (number of fishes in 500 m3) of parrotfish species (± SD) on the reef
flat and slope
t1:2
t1:3 Species Reef flat Reef slope p
t1:4 Cetoscarus bicolor 0.8 ± 1.0 0.7 ± 0.8 0.919 Table 3 t3:1
t1:5 Chlorurus gibbus 0.4 ± 0.7 1.1 ± 1.3 0.282 One-way ANOVA performed on feeding and bioerosion rates of seven species of
t1:6 C. sordidus 7.2 ± 5.8 8.3 ± 6.5 0.749 parrotfishes
t1:7 Scarus ferrugineus 2.4 ± 2.1 3.4 ± 0.9 0.329 t3:2
t1:8 S. frenatus 1.7 ± 2.1 1.9 ± 1.5 0.875 Factor SS df F p t3:3
t1:9 S. ghobban 2.8 ± 2.4 1.8 ± 0.9 0.379 Feeding rates Site 1421.92 5 0.448 0.812 t3:4
t1:10 S. niger 4.1 ± 3.4 4.5 ± 3.2 0.820 Bioerosion rates Site 3798.92 5 1.275 0.296 t3:5

Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
jembe.2009.01.019
 ARTICLE IN PRESS
4 M.A. Alwany et al. / Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx

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Fig. 2. Daily feeding patterns of parrotfish species, averaged over all sites, on the Northern Red Sea reefs. Cetoscarus bicolor (SL 40–50 cm), Chlorurus gibbus (SL 40–45 cm), Chlorurus
sordidus (SL 20–25 cm), Scarus ferrugineus (SL 25–30 cm), S. frenatus (SL 25–35 cm), S. ghobban (SL 35–40 cm), and S. niger (SL 13–25 cm).

238 3.4. Numbers of parrotfish scars on live and dead corals significantly different (p = 0.635). Of the three parrotfishes with high 251
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numbers of scars, two (C. gibbus and S. ghobban) had higher values on 252
239 The average number of parrotfish scars on live corals clearly live coral at the reef slope than the reef flat. The third species, 253
240 differed among the parrotfish species (Table 4 and Fig. 3). Some C. bicolor, had nearly the same values at the two zones. The one-way 254
^
241 species fed on live corals but others definitely avoided them. C. gibbus, ANOVA showed no significant difference between the sites (p = 0.978 255
^ ^
242 C. bicolor and Scarus ghobban belong to the former group on both reef on RF; p = 0.725 on RS), but the number of scars varied highly 256
^ ^
243 zones. C. sordidus and S. niger belong to the latter group. The significantly (p b 0.001 on both RF and RS) between parrotfish species. 257
^
244 remaining species (S. frenatus and S. ferrugineus) only rarely fed on All seven parrotfish species fed on dead coral and hard substrates 258
245 live corals. Compared with the number of scars on dead corals (see that are rich in algae (Table 4). Note that the number of scars on dead 259
246 below), however, live corals did not represent the main food item for corals was generally one order of magnitude higher than on live corals. 260
247 any species. Note that those three species which fed on live corals are S. niger produced the most scars on both zones; C. bicolor had the 261
^ ^
248 also those with higher bioerosion rates (Table 2). They also belong to lowest values. The three main live coral feeders, C. gibbus, C. bicolor 262
^
249 the larger parrotfishes (more than 35 cm SL) studied here. The and S. ghobban, produced the fewest scars on dead coral and hard 263
^
250 difference in scar numbers between reef flat and slope is not substrates. Moreover, the two species (C. sordidus and S. niger) with no 264

Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
jembe.2009.01.019
 ARTICLE IN PRESS
M.A. Alwany et al. / Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx 5

t4:1 Table 4 C. bicolor and C. gibbus fed mainly on live coral. One explanation would 282
Average number of parrotfish scars (±SD 5 min− 1) on live and dead corals of reef flat and be that special or restricted food (high energy content) may not 283
slope
t4:2
always contain all the nutritional components normally found in a 284
t4:3 Species Live corals Dead corals broader variety of food items (low energy content). The remaining 285
t4:4 Reef flat Reef slope Reef flat Reef slope three species (S. ferrugineus, S. frenatus and S. ghobban) had very 286
t4:5 Cetoscarus bicolor 8.4 ± 1.9 8.2 ± 3.2 21.2 ± 9.7 19.8 ± 8.9 similar, intermediate feeding rates and are also relatively similar in 287
t4:6 Chlorurus gibbus 6.4 ± 2.1 8.6 ± 1.4 28.4 ± 13.2 24.8 ± 12.0 size. Beyond species, the sites also played a role in the feeding rates, 288
t4:7 C. sordidus 0 0 74.7 ± 10.8 80.0 ± 11.7 with the within-region differences often being greater than the 289
t4:8 Scarus ferrugineus 0.2 ± 0.5 2.1 ± 4.6 52.3 ± 11.1 55.6 ± 12.8 ^
t4:9
between-region differences. Feeding rates did, however, significantly 290
S. frenatus 0.4 ± 1.0 0.1 ± 0.3 53.1 ± 9.4 55.9 ± 11.4 ^
t4:10 S. ghobban 5.6 ± 1.7 8.9 ± 5.3 45.8 ± 13.6 43.4 ± 14.7 differ between different coral reef habitats. For example, values were 291
t4:11 S. niger 0 0.1 ± 0.3 91.1 ± 12.5 96.8 ± 9.6 highest in the Coral-rich habitat for species feeds on live coral, while 292
^
the dead corals and/or hard substrate habitat had the lowest rates for 293
the same species. 294
265 feeding activity on live coral had high values here. Analysis of variance In contrast to the above differences, the daily feeding patterns of all 295
266 (ANOVA) identified no significant site and reef zone effects (Table 5), species are quite similar, although the large and small species differed 296
267 so the results can therefore be considered to be generally valid for somewhat (Fig. 2). Most species showed a slight peak in activity in the 297
268 Northern Red Sea reefs. afternoon (1400 to 1700 h). This finding confirms for the Red Sea the 298

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results reported by Choat and Clements (1993) and Ferreira et al. 299
^
269 4. Discussion (1998) for other areas. There was also a common and therefore 300
relatively clear trend for a decrease in feeding rate over the course of 301
270 This study on seven parrotfish species provides the first direct the afternoon, with lowest values by the last observation period for all 302
271 estimate of bioerosion rates of this group on Egyptian Red Sea reefs. To species. Choat and Clements (1993) reported that parrotfishes are 303

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272 date, such data has been restricted to the Caribbean and Great Barrier non-selective feeders on complexes of algae, which are ingested with 304
^
273 Reefs. The field observations of foraging behaviour and spatial large amounts of calcareous materials. The greater number of bites on 305
274 distribution (combined with quantifying scar number and size) dead corals and other hard substrates is indirect evidence for a 306
275 revealed considerable differences among parrotfish species and sites preferred algal diet. 307
276 in terms of feeding ecology on Northern Red Sea reefs.

277

278
4.1. Parrotfish feeding activities

The small parrotfishes S. niger and C. sordidus had the highest

^ ^
DP 4.2. Parrotfish bioerosion

Reef eroders include parrotfishes, sea urchins, worms and gastro-

pods, but erosion by the latter two is insignificant (Scoffin et al., 1980).
^
308

309
310
279 feeding rates, whereas the large species C. bicolor and C. gibbus had Parrotfishes are one of the dominant bioeroders on the Great Barrier 311
^
280 low rates. Moreover, the smaller S. niger and C. sordidus fed on dead Reef (Bellwood, 1986, Hutchings, 1986, Kiene, 1988), the Caribbean 312
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281 corals and/or hard substrate (rich in algae), whereas the larger (Gygi, 1975; Garzon-Ferreira and Reyes-Nivia, 2001; Rotjan and Lewis, 313
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Fig. 3. A key consideration in parrotfish bioerosion is to distinguish between feeding on live (A and B, above) versus dead (C and D, below) corals.

Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
jembe.2009.01.019
 ARTICLE IN PRESS
6 M.A. Alwany et al. / Journal of Experimental Marine Biology and Ecology xxx (2009) xxx–xxx

t5:1 Table 5 (1995b) recorded C. gibbus feeding frequently within territories of the 364
Two-way ANOVA performed on abundance and bites on both live and dead corals of striped surgeonfish (Acanthurus lineatus), while the surgeonfish did 365
seven species of parrotfishes ^
permit C. sordidus to feed within the same territories. This also holds 366
t5:2
t5:3 SS df F p true in the Red Sea: C. gibbus feeds within surgeonfish (A. sohal) 367
t5:4 Abundance Site 106.63 5 1.67 0.154 territories. This provides further evidence that C. gibbus mainly feeds 368
t5:5 Reef zone 2.46 1 0.192 0.663 on live coral and not on other substrates, because C. sordidus is a direct 369
t5:6 Site × reef zone 8.79 5 0.137 0.983
competitor of the algal-feeding surgeonfish (e.g. A. sohal), but 370
t5:7 Bites on live corals Site 64.29 5 0.650 0.663 ^
t5:8 Reef zone 22.45 1 1.093 0.290 C. gibbus is not (Alwany et al., 2005). 371
^
t5:9 Site × reef zone 18.61 5 0.188 0.966 Our results provide a number of insights into parrotfish bioerosion. 372
t5:10 Bites on dead corals Site 4126.76 5 1.093 0.372 Importantly, the role of individual species can only be determined by 373
t5:11 Reef zone 39.33 1 0.052 0.820 combining information on abundance, feeding rate, fish activity, food 374
t5:12 Site × reef zone 429.53 5 0.114 0.989
preference and bite volumes. Based on this synthesis we demonstrate 375
an important role of parrotfish species in the Northern Red Sea reefs, 376
particularly of S. ghobban. The next step is to focus on the food 377
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314 2006), French Polynesia (Chazottes et al., 1995) and also in the selection and preferences of parrotfishes for particular live coral 378
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315 Egyptian Red Sea. In the Red Sea, Hassan (1997) reported that species and particular algae species. 379
316 bioerosion was dominated by grazing. While sea urchins, mainly

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317 Diadema setosum, were the main grazers at Aqaba, fishes (parrotfish) Acknowledgements 380
318 dominated at Zabargad Island (Red Sea, Egypt).
319 Recent work by Bellwood (1995a) in the Indo- Pacific and This study was supported by a grant of the ÖAD, Austria, to the first 381
^
320 Bruggemann et al. (1996) in the Caribbean enabled a comparison of author. We thank Prof. M. H. Hanafy and Prof. S. El-Etreby for their 382
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321 parrotfish bioerosion rates among Indo-Pacific, Caribbean and Red Sea suggestions and encouragement in planning this study. This work would 383

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322 reefs. The Caribbean has only one significant bioeroding species not have been possible without the kind assistance of the Department of 384
323 (Sparisoma viride), the Indo-Pacific 16. Furthermore, the latter species Marine Science (Suez Canal University, Egypt) and the Institute of 385
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324 are widespread and relatively abundant (Bellwood, 1995a). Our study Zoology and Limnology (University of Innsbruck, Austria). The authors 386
325 revealed seven bioeroding species in three genera (Chlorurus, Cetos- wish to thank Prof. Joerg Ott, Department of Marine Biology at the 387
326 carus and Scarus). Bellwood (1995a) calculated the annual bioerosion University of Vienna in Austria, for providing working space. [SS] 388
327
328
329
330
rates to be 0.41 m3 y− 1 (1017.7 kg y− 1) for C. gibbus and 0.01 m3 y− 1

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annual rates for the same two species were 0.20 m y (290.3 kg y ;

^ ^
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C. gibbus) and 0.03 m3 y− 1 (42.3 kg y− 1; C. sordidus). This comparison
^
(23.6 kg y− 1) for C. sordidus on the Great Barrier Reef. In our study, the
−1
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DP References

Al-Hussaini, A.H., 1945. The anatomy and histology of the alimentary tract of the coral
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feeding fish Scarus sordidus (Klunz.). Bull. Inst. Egypte 27, 55–69.
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331 indicates that the bioerosion rates of the same parrotfish species differ
Al-Hussaini, A.H., 1947. The feeding habits and the morphology of the alimentary tract 392
332 between different reefs (in the former somewhat lower, in the latter ^
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339 both in the Red Sea and in many regions of the Great Barrier Reef Townsville, Qd. 402
340 (Bellwood, 1995a, 1996) due to its high bite volume (Table 2). Relative Bellwood, D.R., 1995a. A direct estimate of bioerosion by two parrotfish species Chlor- 403
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342 in the Red Sea is small. On the other hand, despite its smaller bite Bellwood, D.R., 1995b. Carbonate transport and within-reef patterns of bioerosion and 406
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348 of the coral colonies and thus retard their growth (Frydl, 1979). Bruggemann, J.H., Kuyper, M.W., Breeman, A.M., 1994b. Comparative analysis of 416
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349 Generally, both the grazing scars and bioerosion rates are thought to Sparisoma viride (Scaridae). Mar. Ecol. Prog. Ser. 112, 51–66. 418
350 increase strongly with fish species size (Bruggemann et al., 1996). In the Bruggemann, J.H., van Kessel, A.M., van Rooij, Breeman, A.M., 1996. Bioerosion and 419
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implications of fish size, feeding mode and habitat use. Mar. Ecol. Prog. Ser. 134, 59–71. 421
352 with fish size. We found that three parrotfish species also fed on live Chazottes, V., Le Campion-Alsumard, T., Peyrot-Clausade, M., 1995. Bioerosion rates on 422
353 coral (C. gibbus, C. bicolor and S. ghobban) in both reef zones. In contrast, ^ ^
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357 two largest parrotfish species had the largest bite volumes and fed influence of spatial variation. Mar. Biol. 89, 221–234. 428
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Please cite this article as: Alwany, M.A., et al., Parrotfish bioerosion on Egyptian Red Sea reefs, J. Exp. Mar. Biol. Ecol. (2009), doi:10.1016/j.
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