Biology of slugs (Agriolimacidae and Arionidae: Mollusca) in New Zealand hill country pastures.

Life cycles of the slugs Deroceras reticulatum and Arion intermedius were studied over a 2.5-year period in a sheep-grazed pasture in the Kaimai Range, New Zealand. D. reticulatum approximated a bivoltine phenology, with intervals between consecutive generations ranging from 4 to 7 months and maximum life span from 8 to 12 months. Egg laying occurred in autumn and spring-early summer. Periods of egg hatching were followed by a phase of juvenile growth characterised by linear increase in the logarithm of population mean body weights. This was followed by a phase of reproductive activity. A. intermedius had an annual life cycle. Eggs laid during late summer and autumn hatched during autumn-winter. Immature slugs were characterised by a period of low growth rate during winter followed by a period of rapid growth culminating in reproductive maturity. In both species, the phase of rapid growth rate was associated with enlargement of the hermaphrodite gland. The size of the hermaphrodite gland relative to body weight reached a maximum during the spermatozoon stage, generally coincident with maximum body weight. During the reproductive period of D. reticulatum and A. intermedius the body weight remained relatively stable but the hermaphrodite gland became progressively smaller as the slugs approached the post-reproductive stage. The albumen gland attained maximum weight at the oocyte stage of the hermaphrodite gland, at the onset of reproductive activity. The pasture, strongly dominated by grasses, was heterogenous in slope, vegetation and treading/grazing influences from sheep due to stratification of the habitat into contour tracks, associated with sheep movement around the slope contours, and intertrack areas. High population densities of D. reticulatum were associated with the track edges (kerbs) and the upper intertrack slopes, while densities of A. intermedius were highest for the intertrack slopes. Low densities of both species occurred on the tracks and at sheep campsites. Analysis of cohort life tables indicated that these dispersion patterns were important in the dynamics of the populations. The stage mortality contributing most to the variance in reproductive populations of both species was that between hatching and onset of reproduction; most of the mortality occurred shortly after hatching on south-facing slopes, but tended to occur later in the life cycle on north slopes. For D. reticulatum this mortality was inversely related to initial density and associated, at least in part, with predation by carabids and birds and with treading by sheep. Density relationships, both at whole plot and plot stratum level, indicated that variations in natality had a stabilizing influence on D. reticulatum populations.