The Tidal Rhythm of Emergence, and the Seasonal Variation of This Synchrony, in an Intertidal Midge

The emergence of an intertial midge was investigated at a site on the coast of the Inland Sea of Japan. The population emerging at this site was drawn from a single species of the genus Clunio, probably C. tsushimensis. Emergence was not synchronized with the day-night cycle, but with the tidal cycle. Moreover, the pattern of synchrony changed with season. A bimodal phase appeared in midwinter; but the pattern of synchrony shifted gradually, during January and February, from morning low tides to afternoon low tides, and a unimodal phase appeared in March. This pattern--i.e., synchrony with afternoon low tides--lasted until early October. In mid-October, the synchrony shifted to the morning low tides. Only a brief bimodal phase appeared in autumn. The phase modality was clearly correlated with the height of tides; i.e., when the low waters in a day were very different in height, emergence was synchronized only with the lower one (April to December). During January and February, the higher low tide, as well as the lower low tide, recedes considerably. The exposure of the larval habitat at the higher low tide may stimulate emergence, resulting in bimodal phases in midwinter. But the unimodal pattern in March cannot be accounted for by a simple synchrony with lower low tide, or with exposure of the larval habitat to the air; the day-night cycle not only would be one of the zeitgebers of the tidal rhythm in every season, but also must participate in the expression of the unimodal phase in spring. Furthermore, the number of midges that emerged each day fluctuated with a semilunar cycle with the season. The phase of this rhythm would be shifted by water temperature.