Spawning aggregation of white-streaked grouper Epinephelusongus: spatial distribution and annual variation in the fish density within a spawning ground

White-streaked grouper (Epinephelus ongus) is an important fisheries target and forms spawning aggregations at particular spawning grounds. The aims of the present study were to investigate the ecological characteristics of annual spawning aggregations such as (1) spatial variations in the density of E. ongus at the spawning ground, (2) the relationship between fish density and environmental variables, (3) inter-annual variations in the spawning aggregation, (4) the proportion of males to females at the spawning ground for several days pre-and post-spawning and (5) the relationship between male density and female density at the protected spawning ground, based on observations over five years at an Okinawan coral reef. Although the protected spawning ground area was large (ca. 2,500 m x 700 m), high density of E. ongus (over 25 individuals per 100 m(2)) was found in a limited area (within c.a. 750 m x 50 m). Current velocity and coverage of rocks had significant positive effects on the spatial distribution of E. ongus at the spawning ground. Inter-annual variation in the degree of aggregation was found and this variation was explained by the annual variation of mean seawater temperature during 40 days before the spawning day. The male-female ratio (male:female) at the spawning ground was ca. 3:1 for three years (May 2012, May 2014 and May 2015) whereas > 13: 1 for one year (May 2013). Significant positive relationships between male density and female density were found at the aggregation sites. It is suggested that E. ongus use aggregation sites with greater current velocity to reduce the risk of egg predation and seawater temperature is one of the main factors that is responsible for determining the degree of aggregation. It is also suggested that females possibly select sites with a greater density of males and this selection behavior might be the reason why females arrived at the spawning ground after the arrival of the males. For effective management of spawning grounds, precise site selection as well as the duration of the protection period are suggested to be key aspects to protect the spawning aggregations of E. ongus, which have been currently achieved at the spawning ground.