3.4 DISCUSSION
My study shows that short exposure to low concentrations of copper has complex latent impacts on E. chloroticus juvenile performance, visible only from 8 d postsettlement,and that it especially affects juvenile resistance to further copper exposure.
3.4.1 LATENT EFFECTS OF LARVAL EXPOSURE TO COPPER ON SETTLERS
Successful settlers were larger and had longer spines relative to body size when they had been exposed to the highest concentration of copper as larvae. However, this positive effect at settlement was cancelled by negative subsequent growth. Settlers that had been exposed to the highest copper concentration both early and late during larval development dramatically decreased in both body size and spine length between 8 d and 25 d post-settlement. As a result, at 25 d post-settlement, settlers that had been in the High copper level groups, were substantially smaller than controls.Juveniles were not individually followed during this experiment and thus a decrease in average size may be due to selective mortality of larger individuals. However, the reduction in average size was large (24%), with very little mortality (less than 5%,corresponding to an average of three individuals per container) suggesting that settlers had actually shrunk. Shrinkage in test diameter of adult sea urchins as the result of food limitation has been reported for some species, including E. chloroticus
(Dix 1972, Levitan 1988, Constable 1993). In the current study, food abundance in settlement containers was not directly measured, however a broad estimation of benthic diatom cover was recorded and no difference was noted between treatments.In addition, benthic coverage appeared more than sufficient for grazing pressure (i.e.well developed brown film with sparse grazing tracks) and almost all settlers had visibly full guts at 25 d post-settlement. Furthermore, growth was not related to settler density, as would be expected if food limitation was a factor. This study provides the first evidence that test shrinkage in urchins might occur as a result of environmental stressors other than food limitation. Furthermore, for these very young juveniles it is a latent effect of larval experience, not their current experience as is the case for food limitation. However, further experimental work following individual settlers would be needed to confirm this pattern.
While there was no difference in settler survival for those from different larval copper concentrations in my experimental setting, growth impairment is likely to result in higher subsequent mortality due to predation in the field. Indeed, young juveniles in many benthic species are thought to be the most at risk of predation (Hunt & Scheibling 1997). Predators of juvenile urchins typically include crabs, demersal fish and sea stars (Hunt & Scheibling 1997, Clemente et al. 2013). Although little is known about predation patterns on small juveniles due to inherent difficulties in studying highly cryptic early life stages, Clemente et al. (2013) showed that the smallest Strongylocentrotus purpuratus juveniles tested (5 - 14 mm) were more than three times more likely to be eaten by crabs than larger juveniles, regardless of predator size. It has been suggested that juvenile urchins may reach an threshold size at which the predation rate dramatically decreases (Menge & Sutherland 1976) or at which they may start feeding on macroalgae thus substantially increasing their growth rate (Rowley 1990). Any delay in reaching this escape size would therefore have a strong impact on a juvenile’s chance of survival or growth, thus affecting population dynamics and adult abundance (Ebert 1983, Underwood & Fairweather 1989).Furthermore, if the observed negative growth was due to shrinkage, then juveniles may also be in poorer body condition as was observed for shrinking Heliocidaris erythrogramma (Constable 1993), and thus be less likely to survive in the field. Very few studies have followed the impact of copper on long-lived benthic invertebrates for as long after settlement as in this study (6 weeks post-settlement). Indeed most studies on pollutants are typically either short-term assays (days) or end shortly after settlement. However, Ng & Keough (2003) reported a dramatic decrease in survival and growth following larval copper exposure in bryozoans visible only weeks to months post-settlement. In contrast, no carry-over effects were observed in growth or survival of juvenile sponges up to six months after larval copper exposure(Cebrian & Uriz 2007).