The life history of the tropical shad Tenualosa toli from Sarawak: First evidence of protandry in the Clupeiformes?

Sharp declines in catches prompted a detailed study of the commercially and culturally important 'terubok' Tenualosa toli, which lives in the fast-flowing, turbid estuaries and adjacent shallow coastal waters of Sarawak. Its reproduction, diet, age and growth were investigated. An absence of small females and large males, together with histological data showing transitional gonads, suggest that T. toli is a protandrous hermaphrodite. Ageing based on otoliths indicates that individuals may not live more than about two years. Male fish spawn towards the end of their first year, change sex (transitional gonads were recorded in fish from 14 to 31 cm SL) and spawn as females in their second year. Spawning takes place in the middle reaches of estuaries and females deposit all their eggs at once. Fecundity is linearly related to fish length but shows significant seasonal and site variations. Hermaphroditism is discussed in relation to possible environmental and isolating mechanisms. T. toli is a zooplanktivore eating mainly calanoid copepods. Laser-ablation inductively-coupled plasma mass spectroscopy of trace elements across otoliths revealed that the species does not move into full seawater or freshwater, but completes its life cycle in estuarine and adjacent coastal waters. Therefore T. toli populations in each estuary and adjacent coastal waters may be relatively isolated from one another, and hence particularly susceptible to overfishing.