An Analysis of Variation in Catchability of Green Tiger Prawn, Penaeus
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702 Abstract.–A method for assessing An analysis of variation in catchability of variation in catchability of single cohort fisheries based on catch and effort sta- green tiger prawn, Penaeus semisulcatus, tistics was developed. The results of its application to the Kuwait shrimp, in waters off Kuwait Penaeus semisulcatus, fishery showed that catchability declined gradually with progress of fishing in general, but Yimin Ye there was a rise in December or Janu- Hussain M. A. Mohammed ary. The pattern of variation was asso- Mariculture and Fisheries Department ciated with possible schooling behavior Kuwait Institute for Scientific Research at the beginning of a fishing season. P. O. Box 1638, 22017 Salmiya, Kuwait Initially, catchability was high, but, as intensive fishing dispersed the schools, E-mail address (for Y. Ye)): [email protected] or reduced population density, catcha- bility declined. The midwinter increase in catchability was probably due to an inshore spawning migration. There were also substantial interseasonal Most fishery management regula- Shrimp fisheries are often man- variations in catchability because school- tions are based on estimation of re- aged by a minimum spawning stock ing was associated with strong recruit- source abundance and fishing mor- ment and environmental conditions. strategy to prevent overfishing of tality, of which catchability is an stocks. Under such a strategy, essential element. The existing lit- CPUE during a spawning season is erature on population dynamics usually used as an index of spawn- rarely deals directly with catchability ing stock abundance, and fishing is per se. For most production models closed when CPUE falls below a cer- and cohort models that use abun- tain level (Penn, 1984; Morgan, dance indices, catchability is usu- 1989). If catchability is highly vari- ally taken as constant and, subse- able, indices of spawning stocks and quently, catch per unit of effort the underlying stock recruitment (CPUE) from commercial catch and relationship could be obscured and effort statistics is used as an index perhaps lead to erroneous manage- of stock abundance. ment decisions (Hannah, 1995). The assumption of constant catch- Studies on catchability are essen- ability is rarely, if ever, valid within tial for relating stock abundance to a fishing season (Garcia, 1988; observed CPUE. By investigating Atran and Loesch, 1995; Hannah, within-season changes in catcha- 1995; Arreguin-Sanchez, 1996), par- bility, properties of a stock that are ticularly for short-lived species such not apparent when only annual as certain shrimp in Kuwait’s wa- data are examined, can be under- ters. Catchability depends on many stood better. The objective of this factors: sensory capabilities, behav- paper is to develop a method to es- ioral response of the target species timate changes in catchability from (Penn, 1984; Wassenberg and Hill, commercial catch and effort data. 1990), environmental factors in- cluding temperature (Chittelbor- ough, 1970; Morgan, 1974; Hill, Materials and methods 1985) and wind (Ehrich and Gro- 1 ger ), stock area, the relative dis- The model tribution of fish and fishing (Win- ters and Wheeler, 1985; Hannah, The Penaeus semisulcatus fishery in 1995), and stock abundance (Mac- waters off Kuwait has a single ma- Call, 1976, 1990; Murphy, 1977; jor recruitment in June and July, Csirke, 1989). Ricker (1975) stated that variation in catchability is likely to be the greatest source of 1 Ehrich, S., and D. Groger. 1989. Diurnal variation in catchability of several fish spe- Manuscript accepted 25 June 1998. error in models based on CPUE cies in the North Sea. ICES Council Meet- Fish. Bull. 97:702–712 (1999). with constant catchability. ing (CM) 1989/B:35, 10 p. Ye and Mohammed: Analysis of variation in catchability of Penaeus semisulcatus in waters off Kuwait 703 which, depending on the year, may be supplemented dependent of time throughout the season and by a second, late-summer recruitment in August or catchability is assumed to remain constant within a early September (Mohammed et al., 1996). The suc- month. cess of the second recruitment was responsible for Given catch and effort data, the initial population record landings, for example, in the 1988–89 and number (R), natural mortality rate (M), and six 1989–90 seasons (Siddeek et al., 1994). Total land- monthly catchability rates (qj), a total of eight pa- ings of P. semisulcatus in 1994–95 and 1995–96 were rameters can be estimated in theory by a time-se- 699.7 and 925.0 tonnes, respectively, about 14% and ries fitting method of least squares between observed 19% of the 1988–89 season’s P. semisulcatus catch of and predicted catches. There exists, however, a prob- 5022.4 tonnes (Siddeek et al., 1994). lem of confounding the initial stock number, natural Kuwait’s opening date for shrimping has been set mortality, and catchabilities. A high initial popula- at 1 September since 1987 (Gulland, 1989; Siddeek tion number with high natural mortality and low and Abdul-Ghaffar, 1989). The season is closed when catchability rates gives as good a fit as does a low the CPUE for combined species of the industrial fleet initial population number with a low natural mor- falls below 80–120 kg/boat-day (Xu et al., 1995). In tality and high catchability rates. The following three this case, shrimp trawling is a process that depletes measures were taken to solve this problem. a single cohort. 1) We assumed that We start with three basic equations. The first equa- 3 3 tion is the relation between fishing mortality and 1 R = ∑∑Cf′ , fishing effort: t t (4) q1 t==1 t 1 = Fqftjt, (1) where C't = observed catch at time t. where qj = catchability in month j; This assumption says, in effect, that for a given esti- Ft = fishing mortality in time interval t mate of q1, we have an estimate of the initial popula- (5 days per interval); and tion number (R) from the average catch and effort ft = fishing effort in time interval t. over the first three time intervals (Hilborn and Walters, 1992). 2) It was presumed that natural The second relates the population number at time t mortality was available from other sources. 3) Aux- to the initial cohort number and cumulative fishing iliary abundance index was used in the time series and natural mortalities: fitting as follows: ti −−FtM T J ∑ it (2) 2 2 = it=1=1 SS=−[]ln( C ) ln( C′ )+− W[] ln( X ) ln( X ′ ) , NRet , ∑ tt∑ j j(5) t=1 j=1 where R = initial stock number; where C = predicted catch at time t by Equation N = population number at time t; t t 3; i = time interval; and C ' = observed catch at time t; M = natural mortality. t Xj' = observed index of stock number in month j in relation to the beginning The final equation relates catch to current abundance point of a fishing season; and fishing as well as to natural mortalities, and we X = predicted relative stock abundance in assume that fishing mortality acts independently of j the middle of month j, i.e. predicted natural mortality, taking a fraction of F/(F+M) of stock number in month j divided by the the total mortality, so predicted stock number of the middle point of the beginning month; and F −+()FM (3) W = the weight given to the deviation be- CN= ()1− e t , tt+ tween the predicted and the observed FMt stock indices. where Ct = catch in time interval t. This procedure reduced the number of parameters The three equations describe an exponential decay to be estimated to six. Monthly CPUE of the United process of a single cohort with fishing and natural Fisheries of Kuwait (UFK) fleet was used as auxil- mortalitites. Natural morality is assumed to be in- iary information about stock abundance. 704 Fishery Bulletin 97(3), 1999 The data Table 1 The shrimp fishery in Kuwait consists of two sec- Time series catch, effort, and relative abundance index data tors: industrial trawlers and dhow trawlers (for a of the Kuwait P. semisulcatus fishery in the 1994–95 and review see Mathews and Samuel, 1991; Mathews, 1995–96 seasons. 1994; Ye et al., 1996). The main species caught are P. semisulcatus, Metapenaeus affinis, and Parapenae- Catch Effort Abundance 3 opsis stylifera. Penaeus semisulcatus dominates in (10 number) (boat-day) index the central and southern waters, and the other two Time 1994 1995 1994 1995 1994 1995 species are mainly distributed in northern waters. The dhow boats trawl in relatively shallow waters, 5 Sep 2007 3752 266.2 357.7 and their catches are auctioned at local fish markets 10 Sep 2212 3165 316.3 325.4 (Abdul-Ghaffar and Al-Ghunaim, 1994; Ye et al., 15 Sep 1565 3201 239.9 352.8 1.000 1.000 20 Sep 1592 4502 260.0 528.8 1996). Species composition of the dhow catch varies 25 Sep 1704 1868 295.0 232.5 with changes in fishing grounds and with seasonal 30/Sep 1549 1805 282.9 237.1 migration. The monthly percentages of P. semisu- 5 Oct 1589 1707 490.8 386.0 lcatus in the dhow landings change greatly within a 10 Oct 782 1342 252.5 316.8 season, for example, from 29.6 to 87.1% in 1993–94. 15 Oct 925 1126 310.9 276.5 0.621 0.613 Owing to the difficulty in collecting reliable effort 20 Oct 921 1397 321.1 356.3 25 Oct 914 1354 329.3 356.9 data from the dhow sector, effort is usually standard- 30 Oct 724 1051 268.8 285.1 ized in relation to that of the industrial sector in re- 4 Nov 421 42 199.7 210.6 porting Kuwait’s shrimp fishery statistics.