Life history of Lineus viridis (Müller, 1774) (Heteronemertea, Nemertea)

Abstract

Lineus viridis is a common nemertean species of North-Atlantic intertidal sand flats. Its mating behaviour is peculiar insofar as this species is reported to be polyandric. However, detailed information on this topic is lacking. In order to get more data on the reproduction, oogenesis and life history of this species, a population in the Wadden Sea on the Isle of Sylt (North Sea) was studied between 2005 and 2011. We conducted regular surveys, during which we sampled, measured and recorded the sex status of 25–100 individuals at each sampling event; at least three individuals were fixed for histological studies at each sampling date. In addition, animals were kept in the laboratory for 3 years to complement field data on sexual identity. Lineus viridis was found to reproduce annually in several successive year; the females are significantly larger than the males. Oogenesis starts in spring, shortly after the preceding reproductive period, and continues until the end of December. Spermiogenesis starts in late autumn and also ends late in December. During mating, several males are generally found crawling on a single female, which forms a cocoon that encloses both the female and the associated males. Fertilization is internal. While females discharge all of their eggs during a single mating event and lose more than 40% of their wet weight, males only empty a few of their gonads, and are thus able to fertilize more than one female. Our study clearly shows that Lineus viridis is a perennial, iteroparous species with a pronounced sexual size dimorphism. During this long-term study, no evidence for sequential hermaphroditism has been found. The observed polyandric mating system in this species raises further questions regarding mate and sperm competition that deserve additional research.