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(Loligo Vulgaris Reynaudii) in South African Waters: an Overview C.Johann Augustyn and Beatriz A

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(Loligo Vulgaris Reynaudii) in South African Waters: an Overview C.Johann Augustyn and Beatriz A AUGUSTYN AND ROEL: SOUTH AFRICAN CHOKKA SQUID FISHERY CalCOFl Rep., Vol. 39, 1998 FISHERIES BIOLOGY, STOCK ASSESSMENT, AND MANAGEMENT OF THE CHOKKA SQUID (LOLIGO VULGARIS REYNAUDII) IN SOUTH AFRICAN WATERS: AN OVERVIEW C.JOHANN AUGUSTYN AND BEATRIZ A. ROE1 Sea Fisheries Kesearch Institute Private Bag X2 Rogge Bay 801 2 South Afnca augustyn~!tfn.wcape.gov.za ABSTRACT FISHERIES BIOLOGY OF CHOKKA SQUID Extensive studies of biology and life cycle, and the Extensive research has been conducted on chokka application of some stock-assessment techniques to South squid, mostly since the early 1980s. The following as- African chokka squid (Loligo vukaris vcyuaudii) have con- pects have been investigated. tributed toward formulating management approaches for . Morphological and biochemical systematics (Augustyn the species. Efforts to clarify the systematics preceded and Grant 1988; Vecchione and Lipinski 1995) the biological, behavioral, population dynamics, life- 0 Embryological and larval development (Vecchione cycle, and ecological studies. Management measures have and Lipinski 1995; Blackburn et al., in press) progressed from simple ones designed to order the fish- 0 Ageing methods and physiology of statolith deposi- ery and control effort, to a more structured approach tion (Lipinski 1991, 1993; Lipinski and Durholtz that uses a closed season as the main management tool. 1994, 1996; Durholtz et al. 1995, 1997; Gerneke et Recent modeling studies have indicated that the stocks al. 1995; Durholtz and Lipinski 1997; Lipinski et al. are nevertheless under pressure, at a time when there 1997; Lipinski, Durholtz, and Underhill, in press) is a political imperative to allow new entrants into the . Trophic relationships and feeding behavior, feeding fishery. There is consequently a need to introduce new physiology (Lipinski 1987, 1990, 1992; Lipinski methods of management (while maintaining effort con- and David 1990; Sauer and Lipinski 1991; Smale et trol as opposed to catch control), which may ultimately al. 1995) lead to the introduction of an operational management . Direct stock-assessment methods, including trawl and procedure. hydroacoustic surveys (Hatanaka et al. 1983; Uozumi et al. 1984, 1985; Wallace et al. 1984; Augustyn et al. INTRODUCTION 1993; Lipinski, Hampton, et al.. in press) This paper reviews the progress that has been made 0 Stock-assessment modeling and methods (Augustyn in South Africa since the early 1980s in understanding et al. 1993; Roe1 et al., in press) the biology and life cycle of chokka squid (Loligo vul- . Management (Augustyn 1986; Lipinslu 1990; Augustyn garis reynaudiz]. It describes how this knowledge has been et al. 1992, 1994; Sauer 1995a, b) combined with various stock-assessment techniques to 0 Reproductive histology and fecundity (Badenhorst develop a management approach for a species that is one 1974; Sauer and Lipinski 1990; Peredo et al. 1996; of a group considered to be very difficult to manage Melo and Sauer 1997, in press) effectively (Pierce and Guerra 1994). 0 Reproductive biology and spawning and schooling Until the mid-1980s chokka squid was taken almost behavior (Lipinski 1979; Augustyn 1990; Sauer et exclusively as a bycatch of the demersal trawl fishery. al. 1992; Sauer and Smale 1993; Hanlon et al. 1994; A small-boat, entrepreneurial, handline jigging fishery Sauer, Roberts, et al. 1997) was established in 1985 (Augustyn 1986). It grew 0 Migration (Sauer et al. 1994; O’Dor, Andrade, et al. explosively, but was soon brought under licensed con- 1996; O’Dor, Webber, et al. 1996) trol. Today the chokka squid jigging fishery is based on . Short- and long-term environmental effects on avail- a relatively small (mean catch: about 6,000 metric tons), ability and spawning (Sauer et al. 1991; Roberts and but valuable resource (close to R 104 million, or $22 Sauer 1994; Roberts et al. 1996; Roberts, in press) million; Cochrane et al. 1997) which is exploited . Population dynamics and life-cycle studies (Augustyn entirely in South African waters. Catches have fluctu- 1989, 1991; Sauer 1991, 1993; Sauer et al. 1993; ated between approximately 2,700 and 1 1,000 t per year Augustyn et al. 1994; Lipinski 1994; Booth et al. (fig. 1). 1997; Sauer, Augustyn, and Roberts 1997) 71 AUGUSTYN AND ROEL SOUTH AFRICAN CHOKKA SQUID FISHERY CalCOFl Rep., Vol. 39, 1998 12000 1 10000 -I P ElForeign Trawl W SA Trawl Jiggers 85 86 87 88 89 90 91 92 93 94 95 96 97 I Year Figure 1. Annual catches of South African chokka squid (Loligo vulgaris reynaudii) in three sectors, from the inception of the fishery in 1985 to 1997. Systematics lished an index of abundance (Augustyn 1989, 1991). Several years before the jigging fishery developed, Sea The distribution of mature animals and different size Fisheries Research Institute (SFRI) recognized that classes indicated that the main spawning areas were lo- chokka squid was a potential new resource. A fisheries cated farther east than had earlier been suspected. Even- biological research project was launched in the early tually these surveys made it possible to draw a more 1980s. Investigating the systematics became a priority complete picture of the life cycle, including aspects such because it was realized that, if a fishery were to be es- as deepwater spawning and larval distribution. tablished and properly managed, the stock distribution With the onset of a directed jigging fishery on chokka would have to be delimited and its systematic relation- squid, the focus of the research shifted to the southeast ship with its European and West African counterpart, coast, because it was realized that the major spawning L. vulgaris, established. From a study in which morpho- grounds were located there. Research based on scuba logical measurements, meristic counts, and protein gel diving clarified the understanding of chokka squid’s mat- electrophoresis were used, it was determined that the ing and spawning behavior and of its seasonality and South African species was a distinct and isolated one and probable life cycle (Augustyn 1990; Sauer 1991). At that genetic differences were at the subspecific, rather the same time, the first steps were taken toward using than specific level. On that basis, the two species were statolith daily growth rings to determine the age of squid renamed L. vulgaris vulgaris and L. vulgaris reynaudii and the basis of ring deposition in statoliths (Lipihski (Augustyn and Grant 1988). and Durholtz 1994, 3 996). In the late 1980s a Squid Workmg Group at the SFRI Biological Studies in Cape Town was established, and cooperation with the Initially the trawler bycatch was sampled to collect squid industry began to lend new impetus to the re- basic biological information in an attempt to understand search. New work, includmg migration and tagging stud- development, growth, population structure, maturation ies (Sauer et al. 1994), acoustic studes, and more advanced cycle, and ecology. Later, the spawning grounds in bays behavioral studies using acoustic tags (Sauer, Roberts, et along the southern coast of South Africa were investi- al. 1997), as well as a comprehensive environmental re- gated (Augustyn 1989). The main spawning sites were search project made further contributions to under- then thought to be located in False Bay, near Cape Town. standing the population dynamics of chokka squid. In the early 1980s, the distribution and relative abun- Our current understanding of the life cycle of this dance, as well as basic biology and ecology were directly squid is summarized by the diagram in figure 2. The studied for the first time on the shelf, during three joint species is relatively short-lived, apparently not exceed- Japanese-South African surveys of the South Coast Shelf ing a life span of 18 months. The population is usually (Hatanaka et al. 1983; Uozumi et al. 1984, 1985). These made up of at least two, sometimes three, major cohorts. were soon followed with regular stratified random sam- Spawning usually peaks in spring and early summer, with pling surveys by a new South African research vessel. a variable smaller peak in autumn or winter (Augustyn The surveys encompassed the whole coast and estab- 1989, 1990; Sauer 1991). A large part of the population 72 AUGUSTYN AND ROEL SOUTH AFRICAN CHOKKA SQUID FISHERY CalCOFl Rep., Vol. 39, 1998 s I SOUTH AFRICA .. CAPE PROVlNCE 34" LC-z.--- - _-*' , .--_-- f- Feeding migrations &Adult spawning migrations +....- Planktonic drifl Adult longshore movement - during spawning Season @ spawning grounds I' H = Hatchlings -'-, Occasional deepwater J = Juveniles C8#- spawning H S = Subadults \ A = Adults 1 I I I t I I I 1 t 1a" 20" 22" 24" 26" E Figure 2 Conceptual diagram of the life cycle of chokka squid (Lollgo vulgaris reynaud!!) in its main area of distribution off the south coast of South Africa TCNP = Tsitsikamma Coastal National Park migrates in waves as mature animals to the southeast signaling with distinct body patterns plays an integral role coast, where most spawn in the shallow (2G50 m) spawn- in spawning and other behavior (see fig. 4), and it is thought ing grounds (Augustyn 1989, 1990; Sauer 1991). These that turbid conditions interfere with this behavior. variations are reflected in the catches, as shown in fig- Different types of shoaling behavior have been iden- ure 3. Availability on the spawning grounds appears to tified on the spawning grounds. Spawning squid often be linked to cool, clear, upwelled water, a seasonal fea- form mushroom-shaped aggregations (Sauer et al. 1992). ture at the capes on the southeast coast, which results The structure of these aggregations and the movement from easterly wind forcing (Sauer et al. 1991). When around them has been elucidated by acoustic tagging waters are warmer than about 21°C or become turbid, studies (Sauer, Roberts, et al. 1997). Several other types the squid tend to spawn in deeper water, often deeper of aggregations have also been observed and are char- than 100 m.
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