ANNUAL AND LUNAR REPRODUCTIVE CYCLES IN LITTORINA SAXATILIS (OLIVI) AND DIFFERENCES BETWEEN BREEDING IN THE MARINE FIRTH OF FORTH AND THE FORTH ESTUARY

Gonads were most commonly reduced in October and fullest inMar-Jun with more pronounced seasonal differences in the estuarythan in the firth. Brood-pouches accordingly were fullest inspring-early summer and emptiest in autumn-early winter. Eggsand early embryos predominated in brood-pouches of early springbut were few in Aug-Oct whereas well-developed juveniles werefew in early spring, increased in spring-summer and came topredominate in Jun-Nov. Brood-pouch loads varied with size offemale but, for a standard 10 mm female, were significantlylower in the 2 estuarine sites than in the firth. Changes inmean brood-pouch count varied only 2.5-fold during the yearin the firth but 4.10-fold in the estuary. In vitro development of eggs to shelled juveniles with the velumresorbed took some 68 d at 10°C and 32%. Eggs and late brood-pouchjuveniles were significantly heavier in the estuary than inthe firth (dry weight, ash-free dry weight (AFDW), ash weight).AFDW changed little during development while ash weight (mostlyshell) increased 3.5-4.4-fold. In a laboratory tide tank, femalesreleased 60 d means of <1.3 juveniles d^–1; more inmid- and late year than in Jan-Mar, more from upshore firthfemales than from low shore or estuary females, more when heldat high tide-tank levels than at lower levels, and generallyat rates comparable to release in containers held at the siteof origin. Egg production in laboratory females did not keeppace with release of young and, at the end of trials, brood-pouchcounts were mostly lower than in contemporary free winkles.Females that had released most (e.g. at high tank levels) containedleast brood-pouch embryos and young subsequently. Release of young in the tide tank was markedly faster at timesnear new moon (often reaching or exceeding 2 female^–1d^–1) than near full moon (often <1 female^–1 d^–1).This lunar fluctuation was clearest in fast-releasing sets heldat high- or mid-tide tank levels. It was not observed in Jan-Mar(when release from most females was slow) nor in females atlow tank levels. (Received 3 September 1990; accepted 8 November 1990)