<I>Plectropomus Leopardus</I>
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BULLETIN OF MARINE SCIENCE. 54(1): 332-342. 1994 PRELIMINARY OBSERVATIONS ON THE SPAWNING BEHAVIOR OF CORAL TROUT, PLECTROPOMUS LEOPARD US (PISCES: SERRANIDAE), ON THE GREAT BARRIER REEF Melita A. Samoilys and Lyle C. Squire ABSTRACT The spawning behavior of the coral trout, Plectropornus leopardus. was studied from August to December 1990 at Scott Reef on the northern Great Barrier Reef. A spawning aggregation was located in an area of approximately 1,700 m2, in which coral trout density was monitored using visual census surveys. Trout aggregated in large numbers in October, with a density increase of up to 12.5-fold above the "normal" density (4 fish' 1,000 m-2): the density recorded outside the aggregation period. Trout numbers began increasing on the full moon in October, peaked over the new moon (44 fish' 1,000 m -2), and dropped rapidly after the first quarter. A smaller spawning aggregation (13 fish· 1,000 m - ') was detected during the following new moon. Distinct courtship coloration and displays were observed in males. Courtship displays occurred at all times of the day. Towards dusk small numbers of males established territories in which they courted and spawned with females. Trout spawned in pairs, exhibiting a rapid rush towards the surface, presumably to release gametes. Spawning was only observed in a 22-min period on sunset. We discuss the contribution of location, timing, and behavioral characteristics of spawning aggregations to spawning success. Species of coral trout (Plectropomus spp.), contribute one of the most important fisheries on the Great Barrier Reef (Craik, 1981; McPherson, 1989; Gwynne, 1990; Trainor, 1991). The combined commercial and recreational reef fish catch is over 7,500 tons annually (Trainor, 1991; Blarney and Hundloe, unpubl. data), of which coral trout comprise at least 2,300 tons. Despite this considerable catch, relatively little is known about the life history of coral trout (but see Goeden, 1978; Ferreira, 1993; Ferreira and Russ, in press), especially about the replenish- ment process from spawning to recruitment. This paper is one of a trio aimed to redress this lack of information. We present the first scientific evidence of spawning aggregations of coral trout in Great Barrier Reef (GBR) waters, and describe the behavior of fish in these aggregations. Doherty et al. (1994) report on coral trout recruitment monitored over the same period; Rimmer et al. (1994) discuss in vitro fertilization of eggs obtained from trout in spawning aggregations. Many reef fish species have a marked seasonal reproductive period; spawning may occur for only a few weeks of the year (Thresher, 1984). Some species, including those of the Serranidae (groupers), aggregate in large numbers at specific sites during their spawning season (Johannes, 1980, 1981; Thresher, 1984; Colin et al., 1987; Shapiro, 1987), presumably to facilitate spawning success. The oc- currence of aggregations of commercial food fish is of significant management importance (Johannes, 1978; Parrack and Huntsman, 1982; Sadovy, in press) because species that aggregate are easy targets for fishers. For example, grouper stocks have been depleted through heavy harvesting of spawning aggregations in Palau (Johannes, 1981), and several Nassau grouper aggregations have disap- peared in the western central Atlantic (Colin, 1992; Sadovy, in press). The com- mercial and recreational fishing communities in Queensland have referred to seasonal spawning aggregations of coral trout for some years, and recently, concern of overfishing spawning aggregations has been expressed by fishers in the northern GBR region. This study focussed on Plectropomus leopardus, the most common of the Dve Australian species of coral trout (Randall and Hoese, 1986), which occurs through- 332 SAMOILYS AND SQUIRE: SPAWNING BEHAVIOR OF CORAL TROUT 333 out the GBR. Plectropomus leopardus is a protogynous hermaphrodite (Goeden, 1978), and usually sexes cannot be differentiated externally. However, we dem- onstrate that male coral trout can be identified by the specific color and behavioral patterns they exhibit when breeding (Rimmer et a1., 1994). While working as a commercial fisherman, one of us (LCS) fished numerous Plectropomus spp. aggregations on the northern GBR. The aggregations were always found during the months of September to February; specifically, September to November for P. leopardus in the Cairns area. This corresponds to "spring- early summer" on the Queensland coast, or the onset of the lighter north-easterly winds. Queensland's climate is characterized by a two season year: the moderate to strong south-east trade winds from March to August ("autumn" and "winter"); and the light north-easterlies from September to February ("spring" and "sum- mer"). Fishers' observations of P. leopardus aggregations in September-November coincide with the months of peak reproductive activity of coral trout off Cairns, as determined from high gonad indices (McPherson et al., 1988). Our study spanned this season and was based at one of the areas identified by LCS as an aggregation site, on a reef close to Cairns. Our first objective was to confirm the seasonal aggregation of mature coral trout through systematic surveys of density. These recorded the formation, persistence and dissolution of at least one major aggregation. Our second objective was to observe and document actual spawning thus confirming the aggregations as spawn- ing events. METHODS Study Site. - The area selected for monitoring was located on the reef slope at Scott Reef (Fig. I), where one of us (LCS) had regularly observed apparent seasonal aggregations over the past 12 years. After preliminary observations, an area of 1,700 m2 (hereafter called the "aggregation site") was mapped and reproduced on underwater paper. These sheets were used as proformas to standardise observations collected during censuses. The site was located on an extensive submerged patch reef, separated from the main reef, and consisted of an upper plateau area submerged at approximately 6-8 m depth, with a steep wall to seaward, dropping to a gently sloping sandy bottom at approximately 17-25 m depth. Moderate to strong currents were a regular feature of the area particularly on the full and new moons. Censuses. - The numbers and sizes (fork length, FL) of coral trout on the aggregation site were counted by a standardised underwater visual census (UYC) using SCUBA. Each census was conducted by one trained observer, although different censuses were done by one of three persons. Training involved estimating fish size using wooden models (Bell et aI., 1985; Samoilys and Carlos, 1992). The observer adopted a fixed search pattern following a route swum in a set time of 25 min. This took the observer around the site at a slow but steady pace. The observer searched a path width of approximately 10 m, with some minor variation caused by visibility. The route was designed to allow the observer to view the whole aggregation site by the end of the census giving a total count rather than a transect count. The observer swam at approximately 1.5 m off the bottom, starting along the shallow upper plateau at 6-8 m and then returned along the wall at about 15 m depth, ensuring no overlap in search path. Since trout move, there may have been inaccuracy in the counts (both over or under-estimation) associated with taking 25 min to complete the count (Watson et aI., in press) but any bias should have been consistent among censuses. Trout numbers were recorded by noting the location and size (in 5 em size classes) of each fish and marking them on the topographical map. Fish sex was recorded by noting courtship colors and display behavior, which is only exhibited by males, and noting females as those individuals which males courted. Sex was confirmed by spearing and dissecting selected fish (Rimmer et at., 1994). The majority ofUYC counts were initiated 50-35 min prior to sunset (range 80-15 min, N = 25), and terminated 10-25 min before sunset. Another set of UYC counts (N = II) were conducted during the day, between 8:30 and 15:30 AEST to determine the stability of aggregations. A total of 36 UYC counts were completed between 21 Aug 1990 and 13 Dec 1990. These were spaced over 25 separate days, with at least one census per week, except for the first week of September. For the first nine trips to Scott Reef (to 10 Oct) and on two subsequent trips (17 and 19 Oct), counts were carried out both during the day and at dusk. Thereafter, counts were only carried out at dusk. 334 BULLETIN OF MARINE SCIENCE, VOL 54, NO. I, 1994 1200E Michaelmas Reef N Arlington Ree F ,,,1 Australia Green [s Ia nd Moore .. Reef Elford Fitzroy . Ree F [sl A nd Sudbury ' Reef 17°S o Kilometres 20 .•..-~ ' - ..... Scott o---Nautical 10 i \, " Reef miles Figure [. Location of study site at Scott Reef south of Cairns, northern Great Barrier Reef. On each census water temperature was measured with a handheld mercury thermometer at 15 m depth. An approximate estimate of visibility was made using a fibreglass tape measure to confirm conditions were appropriate for visual surveys. Neighboring areas on Scott Reef, up to 2 km in all directions from the spawning site, were searched on snorkel during daylight hours for additional aggregations. This included the reef edge (slope) and separated reef patches, Behavioral Observations. - Behavioral observations were started on 17 Oct, the 19th day of censusing. Following completion of the count, the observer remained on the site to watch for spawning activity, Observations were conducted from a stationary position in an area with the most trout. These ob- servations were carried out for approximately 30 min or until darkness. Gonad Maturity. -On several visits to Scott Reef, coral trout were captured on spear or line from a second aggregation site, located 1 km from the study site.