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present in high numbers around sewage outfalls (Anger, 1977) and other organic discharges (see Pearson and Rosenberg, 1978) and, therefore, is not likely to decline in abundance due to the reducing sediments under weed mats. Other studies have suggested that the low abundances of P. elegans under higher weed mat densities have been an indirect response: M. fuliginosus increased in the study by Hull (1987) and it was suggested that M. fuliginosus competed with P. elegans. Unfortunately, these studies have not compared the sizes of P. elegans between weed and control plots and, therefore, it is not possible to determine whether adult mortality or recruitment failure was responsible for the negative effect of weed cover on this species abundance. A possible cause of P.elegans decline in the experiment on Drum Sands could have been sediment accumulation rather than the effect of siltation. During the first few days of the experiment, 5-6cm of sediment had accumulated within the weed treatment plots and it is perhaps unlikely that P. elegans individuals were able to cope with such accretion and were therefore buried. This is supported by the size- frequency distributions of P. elegans: although sediment accumulation affected all size classes, the smaller ones were more severely affected, as would have been expected from sediment smothering. Sediment accumulation by mats of Enteromorpha has been reported elsewhere (e.g., Frostick and McCave, 1979) and is probably a common phenomenon on more exposed sandflats. In addition to the detrimental effect on adults, this study indicated that the decrease in abundance of P. elegans in the long term was likely to have been due to a lack of larval recruitment. Bonsdorff (1992) pointed out that the success of the settling stages of benthic infauna are of crucial importance in determining the structure of the community. It was shown in Chapter 3 that while P. elegans is capable of reproducing by both benthic and planktonic larvae, only planktonic larvae were produced by the population at Drum Sands. Size-frequency distributions clearly showed that the decrease in adult density was mostly due to the decrease in the smaller size classes. After 20 weeks, P. elegans size-frequency distributions in weed treatment plots were significantly different from those in the control plots, mostly due to the dramatic reduction of those individuals smaller than 0.33mm wide (5th setiger). Therefore, the P. elegans larval recruitment which occurred at Drum Sands during 105

May-June 1997 (see Chapter 3) did not occur with any success in the weed treatment plots of this experiment. It is not possible to suggest the mechanism responsible for the lack of P. elegans recruitment in the weed treatment plots in this study. Although reduced water velocity in weed-affected areas may facilitate planktonic larval settlement, larvae may be 'filtered out' by the weed (Olafsson, 1988) preventing them reaching the sediment surface. However, many studies (see Raffaelli et al., 1999) have suggested that reduced P. elegans recruitment to weed-affected areas results from decreased juvenile survival in the more reduced sediments and/or smothering due to siltation. Ragnarsson (1996) found that weed significantly reduced P. elegans colonisation of azoic sediments, however, the present study on Drum Sands is the first experimental study which has reported that macroalgal cover has a negative effect on polychaete recruitment to ambient sediments. The densities of the two bivalves, C. edule and M. balthica, in this study were not significantly affected by weed cover. Everett (1994) found that M. balthica numbers decreased in weed plots compared to algal-free plots and suggested that tellinid bivalves are expected to suffer from the physical barrier formed by algae between the sediment surface and the overlying water column. The results obtained by the algal- removal experiment of Cha (in prep.) supported this. In contrast, Hull (1987) reported greater numbers of M. balthica under his experimentally-implanted algal-covered plots compared to unmanipulated controls. Everett (1994) suggested that the contrasting results obtained by his experiment and the one by Hull (1987) could have been due to two factors. Firstly, Hull's method of weed attachment resulted in an unnatural algal-sediment interface. However, this cannot be the reason for the lack of a negative result in the present study because sediment accumulation resulted in the algae being held in place below the sediment surface, similar to natural weed mats. Secondly, Everett (1994) suggested that the Enteromorpha spp. Hull (1987) used, in contrast to the Ulva spp. in Everett's experiment, being filamentous rather than laminar, may not have formed a barrier between the infauna and the water column to the same extent. However, in Hull's study and the present one on Drum Sands, a significant decrease in the redox profiles occurred under the weed mats leading to 106

present in high numbers around sewage outfalls (Anger, 1977) and other organic<br />

discharges (see Pearson and Rosenberg, 1978) and, therefore, is not likely to decline<br />

in abundance due to the reducing sediments under weed mats. Other studies have<br />

suggested that the low abundances of P. elegans under higher weed mat densities have<br />

been an indirect response: M. fuliginosus increased in the study by Hull (1987) and it<br />

was suggested that M. fuliginosus competed with P. elegans. Unfortunately, these<br />

studies have not compared the sizes of P. elegans between weed and control plots and,<br />

therefore, it is not possible to determine whether adult mortality or recruitment failure<br />

was responsible for the negative effect of weed cover on this species abundance.<br />

A possible cause of P.elegans decline in the experiment on Drum Sands could have<br />

been sediment accumulation rather than the effect of siltation. During the first few<br />

days of the experiment, 5-6cm of sediment had accumulated within the weed<br />

treatment plots and it is perhaps unlikely that P. elegans individuals were able to cope<br />

with such accretion and were therefore buried. This is supported by the size-<br />

frequency distributions of P. elegans: although sediment accumulation affected all<br />

size classes, the smaller ones were more severely affected, as would have been<br />

expected from sediment smothering. Sediment accumulation by mats of<br />

Enteromorpha has been reported elsewhere (e.g., Frostick and McCave, 1979) and is<br />

probably a common phenomenon on more exposed sandflats.<br />

In addition to the detrimental effect on adults, this study indicated that the decrease in<br />

abundance of P. elegans in the long term was likely to have been due to a lack of<br />

larval recruitment. Bonsdorff (1992) pointed out that the success of the settling stages<br />

of benthic infauna are of crucial importance in determining the structure of the<br />

community. It was shown in Chapter 3 that while P. elegans is capable of<br />

reproducing by both benthic and planktonic larvae, only planktonic larvae were<br />

produced by the population at Drum Sands. Size-frequency distributions clearly<br />

showed that the decrease in adult density was mostly due to the decrease in the smaller<br />

size classes. After 20 weeks, P. elegans size-frequency distributions in weed<br />

treatment plots were significantly different from those in the control plots, mostly due<br />

to the dramatic reduction of those individuals smaller than 0.33mm wide (5th setiger).<br />

Therefore, the P. elegans larval recruitment which occurred at Drum Sands during<br />

105

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