Stock Assessment of Two Parrotfish, Hipposcarus harid and Scarus ferrugineus in Jeddah, Saudi Arabia
Stock Assessment of Two Parrotfish, Hipposcarus harid and Scarus ferrugineus in Jeddah, Saudi Arabia
Ahmad Osman Mal1 and Mohamed Hosny Gabr1,2,*
1Marine Biology Department, Faculty of Marine Science, King Abdulaziz University, P.O. Box 80207, Jeddah 21589 Saudi Arabia
2National Institute of Oceanography and Fisheries, Suez, Egypt
ABSTRACT
The current stock status of two Parrotfish, Hipposcarus harid and Scarus ferrugineus in Jeddah was assessed. Scales were used for age determination and back calculations of lengthatages. The growth parameters were estimated to be: the asymptotic length L∞ = 54.044 cm, the growth coefficient K = 0.168 year1, and age at zero length to = 0.707 year for H. harid and L∞ = 51.238 cm, K = 0.170 year1, and to = 0.889 year for S. ferrugineus. The total ‘Z’, natural ‘M’, fishing ‘F’ mortality coefficients and current exploitation ‘Ecur’ were 0.96, 0.294, 0.670 year1, and 0.69 year1, respectively for H. harid and 0.77, 0.299, 0.471 year1, and 0.61 year1 respectively for S. ferrugineus. The maximum yield per recruit at the current fishing mortality was 108.1 g for H. harid and 137.6 g for S. ferrugineus. The biological reference points: Fmax = 0.401 year1 and F0.1 = 0.245 year1 for H. harid and Fmax = 0.434 year1 and F0.1 = 0.307 year1 for S. ferrugineus were lower than the current fishing mortality, reflecting an overexploitation for both species. The current fishing mortality is recommended to be decreased to the target reference point F0.1 = 0.453 year1 for H. harid and F0.1 = 0.466 year1 for S. ferrugineus after increasing the age at first capture to be 3 years for both species.
Article Information
Received 29 July 2017
Revised 12 May 2018
Accepted 06 August 2019
Available online 08 May 2020
Authors’ Contribution
AOM and MHG initiated and designed the study and collected the samples. MHG conducted the experiments and wrote the article. AOM reviewed the article.
Key words
Hipposcarus harid, Scarus ferrugineus, Age determination, Growth parameters, Mortality coefficients, Maximum yield per recruit.
DOI: https://dx.doi.org/10.17582/journal.pjz/20170729030714
* Corresponding author: mhosnyg@yahoo.com
00309923/2020/00051709 $ 9.00/0
Copyright 2020 Zoological Society of Pakistan
Introduction
Hipposcarus harid (Forsskål, 1775) and Scarus ferrugineus (Forsskål, 1775) are two species of Parrotfishes. Ongoing phylogenetic and evolutionary research on parrotfishes indicate that they are either a separate family (Scaridae) under the suborder Labroidei (Bellwood, 1994; Randal, 2007) or a subfamily (Scarinae) under the family Labridae (Westneat and Alfaro, 2005). Parrotfishes are a dominant group of herbivorous species that play an important role in bioerosion and hence affect the benthic communities’ structure on coral reefs (Williams and Hatcher 1983; Russ, 1984; Choat and Bellwood, 1991; Alwany et al., 2009).
Parrotfishes are also important fishery resources in coral reef smallscale artisanal fisheries and caught mainly in gillnet and pot fishing gears. As indicated by Lokrantz et al. (2008), the decline in the biomass and abundance of parrotfishes due to overfishing may have severe negative impacts on the coral reefs dynamics and regeneration. In Saudi Arabia, parrotfishes represent an important part of the catch of coral reef fisheries which are abundant along the Red Sea coast. The average annual catch of parrotfishes from the Red Sea coast of Saudi Arabia during the period from 2008–2016 was 364 tones (FAO, 2018). The two parrotfish species Hipposcarus harid and Scarus ferrugineus are a major component of parrotfish catch from Jeddah fisheries.
The growth rates and longevity in some scarid species have been reported in previous studies based on alternating light and dark bands in hard structures (Warner and Downs, 1977; Russ and St. John, 1988; Clifton, 1995; van Rooij et al., 1995). However, there are only a few studies on the age and growth of the two species in the Red Sea (Ali et al., 2011; Mehanna et al., 2014). There are no previous studies available on the stock assessment for these two parrotfish species in Jeddah fisheries.
The aim of the present study was to estimate age and growth using scales as hard structures, mortalities, length and age at first capture, yield and biomass per recruit of the two parrotfish species in Jeddah fisheries.
Materials and methods
Representative samples of the two parrotfish species Hipposcarus harid and Scarus ferrugineus were collected monthly from the daily fish auction held at the main fish landing site of Jeddah (main fish market) during the period from March 2013 to January 2014. Samples were collected randomly from the landed catch which is harvested using the different fishing gears in Jeddah (Fig. 1), to minimize biases in the sample size distributions produced by differences in gear selectivity (Goodyear, 1995).
For each specimen, the total fish length (L) was measured to the nearest 0.1 cm and total body weight (W) was recorded to the nearest 0.1 g. The power equation: W = a L b was used to describe the length weight relationship, where a is the intercept and b is the slope estimated from the linear regression analysis of the following linearized form of the power equation:
ln W = ln a + b ln L
For age determination, scales from behind the left pectoral fin were collected, cleaned in water then dried and mounted between two microscopic glass slides. The mounted scales were examined under a stereozoom microscope (MEIJI) using a digital video camera connected to a personal computer, where Micrometrics SE Premium software was used to capture and save pictures for scale measurements.
Two linear regression analyses were used to describe the relationship between the body length (L) and scale radius (S) for the two species. The first one is the regression of L on S based on the linear form: L = c + d S (c is the intercept, d is the slope), and the second is the regression of S on L based on the linear form: S = e + f L (e is the intercept, f is the slope). The Lengths corresponding to previous years of life were estimated (backcalculated) using three backcalculation methods (Francis, 1990; Pierce et al., 1996):
FraserLee equation (Lee, 1920):
Li = c + (L  c) (Si / S)
Body proportional hypothesis (BPH):
Li = [(c + dSi) / (c + dS)] L
Scale proportional hypothesis (SPH):
Li = (e/f) + [L + (e/f)] (Si / S)
Where, L is the observed (at capture) fish length, S is the scale radius (at capture), Li is the backcalculated length at the time of annulus i formation, Si is the radius of the annulus i. The mean calculated lengths at ages estimated by the three methods were compared using a oneway analysis of variance (ANOVA) test, applied using ‘Statistix 8.1’ software (Analytical Software, Tallahassee, USA).
The asymptotic length (L∞) and growth coefficient (K) are two parameters of the von Bertalanffy (1938) growth equation (VBGE): Lt = L∞ [1 eK (t t0)]. Both parameters were estimated using the method of Ford (1933) and Walford (1946) fitted to the average backcalculated lengthsatages. The third growth parameter (t0) (supposed age at zero length) was estimated, based on the estimated values of L∞ and K, by rearranging the von Bertalanffy growth equation as described by Sparre and venema (1998): t0 = t + 1/K Loge (1 – Lt / L∞).
The performance index of growth in length ‘ Ǿ ‘ for both species was calculated by the formula suggested by Pauly and Munro (1984): Ǿ = Log K + 2 Log L∞. Because this index is based on length which is rarely lost by fish, it is considered the most precise and flexible index of growth performance which can be used to compare growth performances of wild and cultured fish stocks (Mathews and Samuel, 1990).
The linearized lengthconverted catch curve method of Pauly (1983), implemented in the FiSAT II software (Gayanilo et al., 2005), was used to estimate the instantaneous total mortality coefficient ‘Z’. The natural mortality coefficient ‘M’ was estimated by the equation proposed by Jensen (1996) and modified by Hamel (2015) as follows: M = 1.753 K, where K is the growth coefficient. The difference between the total mortality coefficient ‘Z’ and the natural mortality coefficient ‘M’ was accounted to the fishing mortality coefficient ‘F’. The exploitation ratio ‘E’ was expressed by the ratio between F and Z (E = F/Z).
Three types of fishing gears; gillnets, trammel nets and fish pots are working on all sizes in the fish populations of H. harid and S. ferrugineus in Jeddah fisheries. Based on Crone et al. (2013), since the selectivity of at least one gear type; fish pots, is considered to be sigmoidal (similar to the trawl codend selectivity) (Stewart and Ferrell, 2001; Boutson et al., 2009; Songrak et al., 2013), it is assumed, in the present study, that the gear selectivity for H. harid and S. ferrugineus is asymptotic (sigmoidal) and thus the length at first capture ‘Lc’ (the mean selection length at which 50% of the fish that entered the fishing gear are retained) for both species was estimated using the cumulative method described by Pauly (1984b) implemented in the FiSAT II software. The age ‘ tc ‘ corresponding to Lc was estimated by applying the inverse von bertalanffy equation as follows: tc = t0  1/K Loge (1 – Lc / L∞).
The model of Beverton and Holt (1957) was used to estimate the yield per recruit Y/R and biomass per recruit B/R of H. harid and S. ferrugineus in Jeddah fisheries as follows:
Where, Y/R is yield per recruit, F is the fishing mortality coefficient, M is the natural mortality coefficient, tc is the mean age at first capture, tr is the mean age at recruitment, W∞ is the asymptotic weight, Z is total mortality coefficient, K is the growth coefficient and S equal to the equation: S = e – K ( tc – t0 ). The biomass per recruit (B/R) was estimated by dividing the Y/R by the fishing mortality F.
Two yieldbased biological reference points were determined at two levels of length at first capture Lc: Fmax (the fishing mortality level that produces the maximum yield per recruit) and F0.1 (the fishing mortality level at which the slope or marginal increment of the yield per recruit is 10% of its value at the origin, where E=0). The current fishing mortality level Fcur was matched with these reference points. Fmax is referred to as the ‘Limit’ reference point, while F0.1 is considered as the target reference point (Gabriel and Mace, 1999; Cadima, 2003; Hoggarth et al., 2006).
Results
Fishing gears
Scarid species, as herbivores, are usually caught with gillnets, trammel nets and fish traps (pots). Table I shows the specifications of gillnets and trammel nets used to catch most of the parrotfishes landed in Jeddah fisheries. Two fishermen on a wooden boat (7–9 m length) provided with outboard engine of 25–40 HP usually use some units of gillnets to set in the lagoons and trammel nets to set on the reef flat (mainly during daytime) to target parrotfishes among other reef fishes in the coral reef fisheries of Jeddah.
Age determination and backcalculations
By counting the number of annuli that appeared on the scales, the age in years could be assigned to each specimen. Twelve age groups (011) could be determined for H. harid based on scales reading of 667 specimens ranging from 12.4 – 48.2 cm in total length, while fifteen age groups (014) were observed for S. ferrugineus based on 516 specimens ranging in total length from 13.2–48.5 cm.
The relationship between the fish length and scale radius was found to be linear, as shown in Figure 2, and the model of linear regression of L on S was the best fit for this relationship where the regression standardized residuals plotted against predicted lengths showed no clear pattern but random scattering around zero line (Fig. 2). For backcalculations, the fish length  scale radius relationship could be described by the following two linear equations for each species, based on the linear regression analysis for pooled data (sexes combined):
Table I. Specifications of gillnets and trammel nets used in Jeddah coral reef fisheries.
Gear Item 
Gillnets 
Trammel nets 
Unit length 
35  40 m 
45  50 m 
Net depth 
1.0 – 1.2 m 
0.9 – 1 m 
Number of Units per boat 
4  6 
6  8 
Float line material and diameter 
Polyethylene, 5  6 mm 
Polyethylene, 4  5 mm 
Lead line material and diameter 
Polyethylene, 4  5 mm 
Polyethylene, 4  5 mm 
Number and material of floats per unit 
4550 Cork floats, 25 gf 
6070 Cork floats, 25 gf 
Distance between two floats 
85  90 cm 
60  65 cm 
Number of lead sinkers per unit 
70 – 75 (30 g each) 
110 – 120 (35 g each) 
Distance between two sinkers 
65  70 cm 
35  40 cm 
Inner panel mesh size and twine diameter 
62  88 mm; 0.30.4 mm 
50  62 mm; 0.3 mm 
Outer panel mesh size and twine diameter 
Not present 
150  170 mm; 0.4 mm 
Twine material 
Polyamide, Mono or Multifilament 
Polyamide, monofilament 
Table II. The mean backcalculated lengths at ages estimated by three methods: Fraser Lee, Body proportional hypothesis (BPH) and Scale proportional hypothesis (SPH) for H. harid and S. ferrugineus collected from Jeddah fisheries.
Age 
H. harid 
S. ferrugineus 

FraserLee (pooled) 
BPH 
SPH (pooled) 
FraserLee (pooled) 
BPH 
SPH (pooled) 

Females 
Males 
Pooled 
Females 
Males 
Pooled 

1 
13.83 
14.26 
14.58 
13.82 
12.90 
14.59 
14.81 
14.54 
14.59 
13.89 
2 
20.72 
20.82 
21.20 
20.71 
20.36 
21.09 
21.37 
20.97 
21.09 
20.71 
3 
25.80 
25.82 
25.65 
25.79 
25.52 
25.81 
25.29 
26.10 
25.81 
25.55 
4 
29.72 
29.34 
29.73 
29.71 
29.39 
29.22 
29.16 
29.27 
29.21 
28.93 
5 
32.98 
32.80 
33.24 
32.97 
32.66 
32.21 
32.80 
32.18 
32.21 
31.89 
6 
35.84 
 
35.96 
35.83 
35.38 
35.24 
36.42 
35.11 
35.24 
35.01 
7 
38.50 
 
38.60 
38.49 
38.09 
37.45 
38.65 
37.9 
37.44 
37.15 
8 
41.13 
 
41.20 
41.12 
40.87 
39.39 
40.30 
39.11 
39.38 
39.17 
9 
43.53 
 
43.57 
43.52 
43.36 
41.16 
41.73 
41.00 
41.16 
41.02 
10 
45.30 
 
45.34 
45.29 
45.14 
42.79 
 
42.80 
42.78 
42.68 
11 
46.78 
 
46.79 
46.77 
46.70 
44.30 
 
44.30 
44.29 
44.24 
12 
 
 
 
 
 
45.58 
 
45.59 
45.58 
45.47 
13 
 
 
 
 
 
46.69 
 
46.70 
46.69 
46.62 
14 
 
 
 
 
 
47.60 
 
47.60 
47.59 
47.56 
For H. harid (R2 = 0.917):
L = 2.901 S – 2.175 (regression of L on S)
S = 1.549 + 0.316 L (regression of S on L)
For S. ferrugineus (R2 = 0.930):
L = 2.564 S – 1.726 (regression of L on S)
S = 1.321 + 0.363 L (regression of S on L)
The results listed in Table II and represented in Figure 3 show the mean of the individual backcalculated lengths at ages that estimated by the three backcalculation formulae used in this study. Results of the ANOVA test indicated that there is no statistically significant difference between the mean lengths at ages that estimated by the three backcalculation methods for H. harid (F=0.0034, P=0.9966) and S. ferrugineus (F=0.0023, P=0.9977).
The mean calculated lengths at corresponding ages that estimated by the body proportional hypothesis (BPH) formula for males and females for each species (listed in Table II) were found to be not statistically significantly different (H. harid: F=0.003, P=0.954; S. ferrugineus: F=0.02, P=0.894). So, the growth in length and annual increment were determined from the mean calculated lengths at ages that estimated by the body proportional hypothesis (BPH) formula applied on pooled data (sexes combined) for each species and the results are represented in Figure 4.
The growth equation
Appling the Ford (1933) and Walford (1946) method to the mean lengths at ages estimated by the body proportional hypothesis (BPH) formula applied on pooled data, the growth parameters; L∞, K could be estimated and then the value of t0 was determined and thus the VBGE for describing the growth in length of both species could be written as follows: Lt = 54.0436 [1 e 0.168 (t+0.707)] for H. harid and Lt = 51.238 [1 e 0.170 (t+0.889)] for S. ferrugineus. The von bertalanffy growth curves are shown in Figure 5.
Lengthweight relationship and growth in weight
Total fish lengths and their corresponding weights for H. harid and S. ferrugineus are represented in Figure 6. The length  weight relationship for both species could be described by the nonlinear (power) equations as follows:
For H. harid:
W = 0.026 L 2.81 (R2 = 0.967, n = 73 males)
W = 0.021 L 2.91 (R2 = 0.986, n =594 females)
W = 0.027 L 2.82 (R2 = 0.985, n = 667 both sexes)
For S. ferrugineus:
W = 0.020 L 2.973 (R2 = 0.987, n = 221 males)
W = 0.017 L 3.036 (R2 = 0.985, n = 295 females)
W = 0.020 L 2.970 (R2 = 0.989, n = 516 both sexes)
To check if the growth is isometric (having the value of the exponent ‘b’ equal to 3) or not, the estimated value of ‘b’ for males and females was tested if it was significantly different from 3 or not using the ttest of Pauly (1984a). The results of the ttest indicated that the growth of males and females of H. harid is negatively allometric where the ‘b’ values for both males and females are significantly smaller than the slope value ‘3’ of the isometric growth (for males: t = 3.0852, critical t value = 1.994 for P = 0.05; for females: t = 6.3257, critical t value = 1.965 for P = 0.05). For S. ferrugineus, the growth of males and females was revealed to be isometric, where the b values were found to be not significantly different from the slope ‘3’ of the cubic law of the isometric growth (for males: t = 1.171, critical t value = 1.972 for P = 0.05; for females: t = 1.6449, critical t value = 1.972 for P = 0.05)
Using the obtained equations describing the length  weight relationship for both species, the backcalculated lengths at ages were used to estimate their corresponding weights at ages and thus calculate the growth in weight for both species shown in Figure 7.
Length and age at first capture
Figure 8 shows the results concerning the probability of capture of both species H. harid (A) and S. ferrugineus (B) obtained by the cumulative method described by Pauly (1984b) in the FiSAT II software. The length at 50% probability of capture L50% or Lc and its corresponding age tc was found to be 19.33 cm and 1.93 year for H. harid and 19.30 cm and 1.88 year for S. ferrugineus.
Mortalities and current exploitation
The instantaneous total mortality coefficient Z for H. harid and S. ferrugineus was estimated as the absolute value of the slope of the righthand descending line in Figure 9 (the linearized lengthconverted catch curve as obtained from the FiSAT II software). The value of Z was found to be 0.96 year1 for H. harid and 0.77 year1 for S. ferrugineus. The value of the natural mortality coefficient “M” was estimated as 0.294 year1 for H. harid and 0.299 year1 for S. ferrugineus. The fishing mortality coefficient F was estimated to be 0.670 year1 for H. harid and 0.471 year1 for S. ferrugineus. The current exploitation rate Ecur, which is expressed as the ratio of F/Z, was determined to be 0.69 year1 for H. harid and 0.61 year1 for S. ferrugineus.
Table III. Parameters used to estimate the yield per recruit of H. harid and S. ferrugineus collected from Jeddah fisheries.
Parameter 
H. harid 
S. ferrugineus 
K 
0.168 year1 
0.170 year1 
W∞ 
2078.3 g 
2390.7 g 
t0 
0.7073 year 
0.8894 year 
tc 
1.93 year 
1.88 year 
tc 
2.99 year 
3.04 year 
tr 
0.85 year 
0.77 year 
M 
0.294 year1 
0.299 year1 
Z 
0.96 year1 
0.77 year1 
Fcur 
0.67 year1 
0.471 year1 
F 
Variable 
Variable 
Table IV. The yield per recruit (Y/R) and biomass per recruit (B/R) corresponding to Fcur F0.1 and F max at two values of age at first capture of H. harid and S. ferrugineus in Jeddah fisheries.
Reference point 
H. harid 
S. ferrugineus 

tc = 1.93 year 
tc = 2.99 year 
tc = 1.88 year 
tc = 3.04 year 

F (year1) 
Y/R (g) 
B/R (g) 
F (year1) 
Y/R (g) 
B/R (g) 
F (year1) 
Y/R (g) 
B/R (g) 
F (year1) 
Y/R (g) 
B/R (g) 

F 0.1 
0.245 
107.6 
439.0 
0.453 
126.7 
279.7 
0.307 
134.7 
438.6 
0.466 
153.4 
329.1 
F max 
0.401 
112.5 
280.6 
0.686 
129.0 
188.0 
0.434 
137.7 
317.2 
0.830 
158.1 
190.4 
F cur 
0.670 
108.1 
161.4 
0.670 
129.0 
192.5 
0.471 
137.6 
292.1 
0.471 
153.6 
326.0 
Maximum yield and biomass per recruit
The parameters used to estimate the yield per recruit of H. harid and S. ferrugineus in Jeddah fisheries are given in Table III. The estimated Y/R and B/R as a function of fishing mortality at two values of age at first capture tc = 1.93 and 2.99 year for H. harid and tc =1.88 and 3.04 year for S. ferrugineus are shown in Figure 10.
The yield per recruit and biomass per recruit (in grams) corresponding to the current fishing mortality Fcur relative to those corresponding to the two biological reference points Fmax and F0.1 estimated at the two levels of age at first capture are listed in Table IV. For H. harid, at the current level of fishing mortality Fcur = 0.67 year1 and age at first capture tc = 1.93 year, the yield and biomass per recruit were estimated to be 108.1 and 161.4 g, respectively. For S. ferrugineus, the current level of fishing mortality Fcur = 0.471 year1 and age at first capture tc = 1.88 year, resulted in a yield per recruit of 137.6 g and a biomass per recruit of 292.1 g.
Increasing the age at first capture from 1.93 to 2.99 year (corresponding to 25 cm total length) for H. harid increased the yield and biomass per recruit at the current level of fishing mortality to 129.0 and 192.5 g, respectively. Similarly, when the age at first capture increased from 1.88 to 3.04 year (corresponding to 25 cm total length) for S. ferrugineus, the yield and biomass per recruit at the current level of fishing mortality increased to 153.6 and 326.0 g, respectively.
Discussion
Age determination and backcalculations
In the present study, the scales were used for age determination of H. harid and S. ferrugineus in Jeddah fisheries based on two criteria (Williams and Bedford, 1974): the recognized characteristic pattern shown on the scales (e.g., Fig. 11) and the annual time scale assigned for each pair of alternative opaque and hyaline bands (annulus) as confirmed in previous studies on scarid species (Warner and Downs, 1977; Lou, 1992; Fowler, 1995; Choat et al., 1996; Ali et al., 2011; Mehanna et al., 2014).
In the present study, the relationship between fish length and scale radius showed that the larger the observed fish length (L), the larger the scale radius at capture (S), based on the strong linear relationship: L = 2.901 S – 2.175 for H. harid and L = 2.564 S – 1.726 for S. ferrugineus. For H. harid ranging in total length from 12.4 to 48.2 cm, eleven annuli were laid down on the scales, while for S. ferrugineus ranging in total length from 13.2 to 48.5 cm, fourteen annuli could be observed on the scales.
The backcalculated lengths at ages that are estimated by the FraserLee formula and the body proportional hypothesis (BPH) formula were almost the same for both species. This is because the two methods were based on the same linear regression (L on S), whereas the scale proportional hypothesis (SPH) formula produced slightly lower backcalculated lengths at ages because it is based on a different linear regression (S on L) as indicated by Francis (1990). However, results of the ANOVA test showed that the difference between the backcalculated lengths at ages estimated by the three different methods for the two species (pooled data) was not statistically significant. These results agree well with what is reported by Pierce et al. (1996).
The backcalculated lengths at ages that are estimated by the body proportional hypothesis (BPH) formula, as one of the two proportional hypotheses recommended by Francis (1990) for backcalculations, were considered in the present study to calculate the growth in length and growth parameters for sexes combined (pooled data) since there was no statistically significant difference between the backcalculated lengths at corresponding ages for males and females. Because the two scarid species are protogynous (females change to males in case of deficiency) (Ali et al., 2011; Choat and Robertson, 1975), females predominate in the younger length or age groups while being missed in older groups which are predominated by males.
Table V. Growth parameters of H. harid and S. ferrugineus in the Red Sea estimated by different Authors.
Reference 
H. harid 
S. ferrugineus 

L∞ 
K 
t0 
Ǿ 
L∞ 
K 
t0 
Ǿ 

Present study (Red Sea–Saudi Arabia) 
54.0436 
0.168 
0.7073 
2.69 
51.238 
0.170 
0.8892 
2.65 
Ali et al (2011) (Red Sea Saudi Arabia) 
44.59 
0.17 
1.52 
2.53 
61.4 
0.1 
2.20 
2.60 
Mehanna et al (2014) (Red Sea Egypt) 
57.16 
0.23 
0.69 
2.88* 
 
 
 
 
* Estimated from the available L∞ and K values.
Table VI. Lengthweight relationship parameters of H. harid and S. ferrugineus in the Red Sea estimated by different Authors.
Reference 
H. harid 
S. ferrugineus 

Length range 
a 
b 
r2 
Length range 
a 
b 
r2 

Present study (Red Sea–Saudi Arabia) 
12.4  48.2 TL 
0.027 
2.82 
0.985 
13.2  48.5 TL 
0.020 
2.97 
0.989 
Ali et al (2011) (Red Sea Saudi Arabia) 
15.9 – 31.0 SL 
0.023 
2.99 
 
12.5 – 31.6 SL 
0.019 
3.09 
 
Mehanna et al. (2014) (Red Sea Egypt) 
17.0 – 50.0 TL 
0.018 
2.932 
0.975 
 
 
 
 
TL, total length; SL, standard length.
Growth in length
The growth rates estimated from the average backcalculated lengths at ages for sexes combined (pooled data) indicated that the maximum rate of growth in length was attained during the first year of life for both H. harid (13.82 cm) and S. ferrugineus (14.59 cm). During the second year, the annual increment was reduced to less than half of its value in the first year for H. harid (6.9 cm) and S. ferrugineus (6.5 cm), followed by gradual decrease during the next years. This trend of growth in length is more or less similar to those observed and reported in previous studies on the same two species in the Red Sea (Ali et al., 2011; Mehanna et al., 2014,) and other scarid species (Choat et al., 1996; Taylor and Choat, 2014).
Growth parameters
Considerable variability was found among the von Bertalanffy growth parameters estimated by different authors based on backcalculated lengthsatages, where scales were used for age determination for H. harid and S. ferrugineus in the Red Sea (Table V). This variability might be because of different backcalculated lengths at ages calculated using different backcalculation methods. Collecting scales from different body sites, measuring scales at different angles, and poor representation of all size groups in the samples may lead to a wide variation in the values of the regression analyses parameters used in the backcalculation methods (Carlander, 1982; Hirschhorn and Small, 1987).
For H. harid, the growth parameters estimated in the present study are comparable to those estimated by Mehanna et al (2014). The larger values of the asymptotic length and growth coefficient may be due to the slightly wider length range (17–50 cm) with fewer age groups assigned (8 age groups compared to 12 age groups in the present study).
For S. ferrugineus, Ali et al. (2011) estimated an extremely high asymptotic length (61.4 cm, standard length) with a corresponding very low growth coefficient (0.1 yr1) compared to the maximum observed length they recorded (31.6 cm). In the present study, the estimated asymptotic length (51.2 cm total length) was close to the maximum observed length (48.5 cm) and much smaller than that estimated by Ali et al. (2011).
However, both values of the growth coefficient for H. harid (0.168 year1) and S. ferrugineus (0.1704 year1) indicate that both species are moderate growth species (Branstetter,1987). H. harid has slightly larger growth performance than that of S. ferrugineus due to the larger asymptotic length of the former (Table V).
Lengthweight relationship
From the power equation describing the relationship between fish length and weight, the fish growth can be determined to be isometric or not. When the exponent ‘b’ equals ‘3’ then the growth is isometric. Values of ‘b’ lower than ‘3’ mean negative allometric growth, whereas b values higher than ‘3’ refer to positive allometric growth (Froese, 2006). As indicated in Table VI, the value of the exponent ‘ b ‘ for H. harid was significantly lower than ‘ 3 ‘, reflecting a negative allometric growth for this species in Jeddah fisheries.
Mehanna et al. (2014) predicted a negative allometric growth for the same species having comparable size range and length type (total length) in the Egyptian Red Sea. Ali et al. (2011) used smaller size range and length type (standard length) for the same species in Jeddah fisheries estimated the ‘b’ value to be 2.99 which is not significantly different from the isometric growth (b = 3). Also, Ali et al. (2011) estimated higher ‘b’ value for S. ferrugineus (3.09) compared to that found in the present study (2.97). However, Carlander (1977) showed that using the standardlength measurements results in higher condition factors than that when using measurements of larger length types (fork and total length). This, in addition to the narrow size range, may be the reason behind the larger ‘b’ values obtained by Ali et al. (2011) compared to that obtained in the present study.
Exploitation and maximum yield per recruit
The obtained results of mortality coefficients listed in Table III indicate that the stocks of both species are subject to a fishing mortality that is almost double the amount of natural deaths. To assess the stock status of both species under the current level of fishing mortality, the Beverton and Holt’s model (Beverton and Holt, 1957) was applied to estimate the yield per recruit at various levels of fishing mortality giving the yield per recruit curve for each species (Fig. 10).
For H. harid, it is evident from the results listed in Table IV and shown in Figure 10, that the current fishing mortality rate Fcur = 0.67 year1 is higher than both values of Fmax = 0.401 year1 and F0.1 = 0.245 year1, which means that the stock of this species is currently overexploited. Increasing the age at first capture from 1.93 to 2.99 year increased the yield and biomass per recruit at the current level of fishing mortality from 108.1 and 161.4 g (12.4% of the virgin (unfished) biomass per recruit ‘Bv/R=1305.29 g’) to 129.0 and 192.5 g (14.7% of Bv/R), respectively.
For S. ferrugineus, the current level of fishing mortality Fcur = 0.471 year1 is higher than both values of Fmax = 0.434 year1 and F0.1 = 0.307 year1, reflecting a current overexploitation of the stock of this species in Jeddah fisheries. Similarly, when the age at first capture increased from 1.88 to 3.04 year, the yield and biomass per recruit at the current level of fishing mortality increased from 137.6 and 292.1 g (19.3% of Bv/R=1516.2 g) to 153.6 and 326.0 g (21.5% of Bv/R), respectively.
For both species, the levels of biomass per recruit at the current levels of fishing mortality may be not enough to provide sufficient recruitment to the fishery. Goodyear (1993) indicated that the recruitment process is dependent on the spawning stock biomass per recruit (SSBR). He also reported that the spawning potential ratio SPR = SSBRfished / SSBRunfished, which is proportional to B/Rfished / B/Runfished expressed in the Beverton and Holt’s yield per recruit model, should be between 20–30% to obtain the maximum or near maximum yield per recruit. Thus, we recommend reducing the current level of fishing mortality to that of the target reference point F0.1 = 0.453 year1 after increasing the age at first capture from 1.93 year to 2.99 year to get a biomass per recruit of 279.7 g (21.4 %) for H. harid. For S. ferrugineus, we recommend reducing the fishing mortality from the current level to that of the target reference point F0.1 = 0.466 year1 after increasing the age at first capture from 1.88 year to 3.04 year to get a biomass per recruit of 329.1 g (21.7 %).
However, increasing the age at first capture means improving the fishing gear selectivity to avoid the capture of young immature fish; by increasing the mesh size of the fishing net or/and avoiding the fishing ground having the young immature fish. Gabr and Mal (2016) recommended the use of trammel nets with 62 mm innerpanel mesh size to catch H. harid of a mean selection length of 24.46 cm total length, which is very close to that recommended in the present study to obtain the target yield and biomass per recruit.
Conclusion
Currently, the stocks of both species Hipposcarus harid and Scarus ferrugineus in Jeddah fisheries are overexploited. For both species, the levels of biomass per recruit at the current levels of fishing mortality may be not enough to provide sufficient recruitment to the fishery. Thus, we recommend reducing the current level of fishing mortality to that of the target reference point F0.1 = 0.453 year1 after increasing the age at first capture from 1.93 year to 2.99 year to get a biomass per recruit of 279.7 g (21.4 % of the virgin stock biomass per recruit) for H. harid. For S. ferrugineus, we recommend reducing the fishing mortality from the current level to that of the target reference point F0.1 = 0.466 year1 after increasing the age at first capture from 1.88 year to 3.04 year to get a biomass per recruit of 329.1 g (21.7 % of the virgin stock biomass per recruit).
Acknowledgments
This project was funded by the Deanship of Scientific Research (DSR), at King Abdulaziz University, Jeddah, under Grant No. 461/150/1434. The authors, therefore, acknowledge with thanks DSR for technical and financial support.
Statement of conflict of interest
The authors declare no conflict of interest.
References
Ali, T.E., Osman, A.M., AbdelAziz, S.H. and Bawazeer, F.A., 2011. Growth and longevity of the protogynous parrotfish, Hipposcarus harid, Scarus ferrugineus and Chlorurus sordidus (Teleostei, Scaridae), off the eastern coast of the Red Sea. J. appl. Ichthyol., 27: 840–846. https://doi.org/10.1111/j.14390426.2010.01566.x
Alwany, M.A., Thaler, E. and Stachowitsch, M., 2009. Parrotfish bioerosion on Egyptian Red Sea reefs. J. exp. Mar. Biol. Ecol., 371: 170176. https://doi.org/10.1016/j.jembe.2009.01.019
Bellwood, D.R., 1994. A phylogenetic study of the parrotfishes family Scaridae (Pisces: Labroidei), with a revision of genera. Rec. Aust. Mus. (Suppl.), 20: 186. https://doi.org/10.3853/j.08127387.20.1994.51
Beverton, R.J.H. and Holt, S.J., 1957. On the dynamics of exploited fish populations. Fishery Invest, Ministry of Agriculture of Fish and Food, G.B. London, pp. 533. https://trove.nla.gov.au/work/13338365
Boutson, A., Mahasawasde, C., Mahasawasde, S., Tunkijanukij, S. and Arimoto, T., 2009. Use of escape vents to improve size and species selectivity of collapsible pot for blue swimming crab Portunus pelagicus in Thailand. Fish. Sci., 75: 2533. https://doi.org/10.1007/s125620080010z
Branstetter, S., 1987. Age, growth and reproductive biology of the silky shark, Carcharhinus falciformis, and the scalloped hammerhead, Sphyrna lewini, from the northwestern Gulf of Mexico. Environ. Biol. Fish., 19: 161–173. https://doi.org/10.1007/BF00005346
Cadima, E.L., 2003. Fish stock assessment manual. FAO Fisheries Technical Paper 393, FAO, pp. 161. http://www.fao.org/3/ax8498e.pdf
Carlander, K.D., 1982. Standard intercepts for calculating lengths from scale measurements for some centrarchid and percid fishes. Trans. Am. Fish. Soc., 111: 332336. https://doi.org/10.1577/15488659(1982)111<332:SIFCLF>2.0.CO;2
Choat, J.H. and Robertson, D.R., 1975. Protogynous hermaphroditism in fishes of the family Scaridae. In: Intersexuality in the animal kingdom (ed. R. Reinboth). Springer, Berlin, Heidelberg, pp. 263283. https://doi.org/10.1007/9783642660696_26
Choat, J.H. and Bellwood, D.R., 1991. Reef fishes their history and evolution. In: The ecology of fishes on coral reefs (ed. P.F. Sale). Academic Press, San Diego, CA, pp. 3966. https://doi.org/10.1016/B9780080925516.500088
Choat, J.H., Axe, L.M. and Lou, D.C., 1996. Growth and longevity in fishes of the family Scaridae. Mar. Ecol. Progr. Ser., 145: 33–41. https://doi.org/10.3354/meps145033
Clifton, K.E., 1995. Asynchronous food availability on neighboring Caribbean coral reefs determines seasonal patterns of growth and reproduction for the herbivorous parrotfish Scarus iserti. Mar. Ecol. Prog. Ser., 116: 3946. https://doi.org/10.3354/meps116039
Crone, P., Maunder, M., Valero, J., McDaniel, J. and Semmens, B., 2013. Selectivity: Theory, estimation, and application in fishery stock assessment models. Center for the Advancement of Population Assessment Methodology (CAPAM), Workshop Series Report 1, pp. 46. https://doi.org/10.1016/j.fishres.2014.03.017
FAO, 2018. Fishery and aquaculture statistics. Global Capture Production 19502016 (FishstatJ). FAO Fisheries and Aquaculture Department, Rome. www.fao.org/fishery/statistics/software/fishstatj/en
Ford, E., 1933. An account of the herring investigations conducted at Plymouth during the years from 1924 to 1933. J. Mar. Biol. Assoc. U.K., 19: 305–384. https://doi.org/10.1017/S0025315400055910
Fowler, A.J., 1995. Annulus formation on otoliths of coral reef fisha review. In: Recent developments in fish otolith research (eds. D.H. Secor, J.M. Dean and S.E. Campana). University of South Carolina Press, Columbia, pp. 4563.
Francis, R.I.C.C., 1990. Backcalculation of fish length: a critical review. J. Fish Biol., 36: 883902. https://doi.org/10.1111/j.10958649.1990.tb05636.x
Froese, R., 2006. Cube law, condition factor, and weightlength relationships: History, meta405 analysis and recommendations. J. appl. Ichthyol., 22: 241253. https://doi.org/10.1111/j.14390426.2006.00805.x
Gabr, M.H. and Mal, A.O., 2016. Trammel net sizeselectivity for Hipposcarus harid (Forsskal, 1775) and Lethrinus harak (Forsskal, 1775) in coral reef fisheries of Jeddah, Saudi Arabia. Egyptian J. aquat. Res., 42: 491498. https://doi.org/10.1016/j.ejar.2016.09.005
Gabriel, W.L. and Mace, P.M., 1999. A review of biological reference points in the context of the precautionary approach. Proceedings of the 5th annual NMFS National Stock Assessment Workshop, NOAA Tech. Memo., NMFSF/SPO40. https://www.st.nmfs.noaa.gov/Assets/stock/documents/workshops/nsaw_5/gabriel_.pdf
Gayanilo, Jr. F.C., Sparre, P. and Pauly, D., 2005. User’s guide: FAOICLARM stock assessment tools II (FiSAT II), revised version. FAO Computerized Information Series (Fisheries) No. 8, revised version. FAO, Rome, pp. 168.
Goodyear, C.P., 1993. Spawning stock biomass per recruit in fisheries management: foundation and current use. In: Risk evaluation and biological reference points for fisheries management (eds. S.J. Smith, J.J. Hunt and D. Rivard). Canadian Special Publication of Fisheries and Aquatic Sciences, pp. 6781.
Goodyear, C.P., 1995. Mean size at age: An evaluation of sampling strategies using simulated red grouper data. Trans. Am. Fish. Soc., 124: 746–755. https://doi.org/10.1577/15488659(1995)124<0746:MSAAAE>2.3.CO;2
Hamel, O.S., 2015. A method for calculating a metaanalytical prior for the natural mortality rate using multiple life history correlates. ICES J. Mar. Sci., 72: 62–69. https://doi.org/10.1093/icesjms/fsu131
Hirschhorn, G. and Small, G.J., 1987. Variability in growth parameter estimates from scales of Pacific cod based on scale and area measurements. In: Age and growth of fish (eds. R.C. Summerfelt and G.E. Hall). Iowa State University Press, Ames, pp. 147157.
Hoggarth, D.D., Abeyasekera, S., Arthur, R.I., Beddington, J.R., Burn, R.W., Halls, A.S., Kirkwood, G.P., McAllister, M., Medley, P., Mees, C.C., Parkes, G.B., Pilling, G.M., Wakeford, R.C. and Welcomme, R.L., 2006. Stock Assessment for fishery management – A Framework guide to the stock assessment tools of the fisheries management science programme (FMSP). FAO Fisheries Technical Paper 487, pp. 261. http://www.fao.org/3/aa0486e.pdf
Jensen, A.L., 1996. Beverton and Holt life history invariants result from optimal tradeoff of reproduction and survival. Can. J. Fish. aquat. Sci., 53: 820–822. https://doi.org/10.1139/f95233
Lee, R.M., 1920. A review of the methods of age and growth determination in fishes by means of scales. Fish. Investig. Lond. Ser., 24: 1–32.
Lokrantz, J., Nystrom, M., Thyresson, M. and Johansson, C., 2008. The nonlinear relationship between body size and function in parrotfishes. Coral Reefs, 27: 967974. https://doi.org/10.1007/s0033800803943
Lou, D.C., 1992. Validation of annual growth bands on the otolith of tropical parrotfishes (Scarus schlegeli Bleeker). J. Fish. Biol., 41: 775–790. https://doi.org/10.1111/j.10958649.1992.tb02706.x
Mathews, C.P. and Samuel, M., 1990. Using the growth performance index Ǿ to choose species for aquaculture: an example from Kuwait. Aquabyte, 3: 24.
Mehanna, S.F., AbuElregal, M. and AbdelMaksoud, Y.A., 2014. Age and growth based on the scale readings of the two scarid species Hipposcarus harid and Chlorurus sordidus from Hurgada fishing area, Red Sea, Egypt. Int. J. mar. Sci., 4: 16.
Pauly, D., 1983. Some simple methods for the assessment of tropical fish stocks. FAO Fisheries Technical Paper 234, FAO, Rome, pp. 52.
Pauly, D., 1984a. Fish population dynamics in tropical water: A manual for use with programmable calculators. ICLARM Studies and Reviews 8, International Center for Living Aquatic Resources Management, Manila, Philippines.
Pauly, D., 1984b. Lengthconverted catch curves. A powerful tool for fisheries research in the Tropics (Part 1). ICLARM Fishbyte, 1: 913.
Pauly, D. and Munro, J.I., 1984. Once more on the comparison of growth in fish and invertebrates. FishByte, 2: 2123.
Pierce, C.L., Rasmussen, J.B. and Leggett, W., 1996. Backcalculation of fish length from scales: Empirical comparison of proportional methods. Trans. Am. Fish. Soc., 125: 889918. https://doi.org/10.1577/15488659(1996)125<0889:BCOFLF>2.3.CO;2
Randall, J.E., 2007. Reef and shore fishes of the Hawaiian Islands. University of Hawai‘i Sea Grant College Program, Honolulu. ISBN 9781929054039 https://doi.org/10.1643/00458511(2007)2007[775:RASFOT]2.0.CO;2
Russ, G.R., 1984. Distribution and abundance of herbivorous grazing fishes in the central Great Barrier Reef. 11. Patterns of zonation of midshelf and outershelf reefs. Mar. Ecol. Progr. Ser., 20: 3544. https://doi.org/10.3354/meps020035
Russ, G.R. and St. John, J., 1988. Diets, growth rates and secondary production of herbivorous coral reef fishes. Proceedings of the Sixth International Coral Reef Symposium, pp. 3743.
Songrak, A., Bodhisuwan, W. and ThapanandChaidee, T., 2013. Selectivity of traps for blue swimming crab in Trang province. MAEJO Int. J. Sci. Technol., 7: 3642.
Sparre, P. and Venema, S.C., 1998. Introduction to tropical fish stock assessment, Part 1. Manual. FAO Fisheries Technical Paper. No. 306.1, Rev. 2, pp. 407. ftp://ftp.fao.org/docrep/fao/w5449e/w5449e00.pdf
Stewart, J. and Ferrell, D.J., 2001. Mesh selectivity in the NSW demersal trap fishery. Fish. Res., 59: 379392. https://doi.org/10.1016/S01657836(02)000243
Taylor, B.M. and Choat, J.H., 2014. Comparative demography of commercially important parrotfish species from Micronesia. J. Fish Biol., 84: 383402. https://doi.org/10.1111/jfb.12294
van Rooij, J.M., Bruggemann, J.H., Videler, J.J. and Breeman, A.M., 1995. Plastic growth of the herbivorous reef fish Sparisoma viride—field evidence for a tradeoff between growth and reproduction. Mar. Ecol. Progr. Ser., 122: 93105. https://doi.org/10.3354/meps122093
von Bertalanffy, L., 1938. A quantitative theory of organic growth (inquiries on growth laws 2). Human Biol., 10: 181–213.
Walford, L.A., 1946. A new graphic method of describing the growth of animals. Biol. Bull. Mar. Biol. Lab., 90: 141–147. https://doi.org/10.2307/1538217
Warner, R.R. and Downs, I.F., 1977. Comparative life histories: Growth vs reproduction in normal males and sexchanging hermaphrodites of the striped parrotfish, Scarus croicensis. Proceeding of the 3rd International Coral Reef Symposium, pp. 275281.
Williams, T. and Bedford, B.C., 1974. The use of otoliths for age determination. In: The ageing of fish (ed. T.B. Baginal). Unwin Brothers Ltd., Surrey, pp. 114–123.
Williams, D.B. and Hatcher, A.I., 1983. Structure of fish communities on outer slopes of inshore, midshelf and outer shelf reefs of the Great Barrier Reef. Mar. Ecol. Progr. Ser., 10: 239250. https://doi.org/10.3354/meps010239
Westneat, M.W. and Alfaro, M.E., 2005. Phylogenetic relationships and evolutionary history of the reef fish family Labridae. Mol. Phylogen. Evolut., 36: 370390. https://doi.org/10.1016/j.ympev.2005.02.001
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