Biodiversity and ecosystem functioning: issues of scale and trophic ...

Raffaelli: Biodiversity and ecosystem functioning289the sediment (Raffaelli 2000b), as well as presenting abarrier to the selective feeding of higher trophic levels,especially fish and shorebirds (Raffaelli et al. 1998),and these species may then leave the location altogether(Raffaelli et al. 1999) (Fig. 1). The feedbacks onmacrofaunal diversity will, in turn, alter ammoniumrelease, so that the net effects of macrofaunal speciesrichness on ecosystem functioning will be muchmore complex than is revealed in single trophic levelexperiments.A further reason for adopting a multi-trophic perspectiveis the need to recognise that impacts whichlead to biodiversity loss usually occur across all trophiclevels and that, as Solan et al. (2004) have shown, taxawill differ in their sensitivities to particular loss scenarios(Fig. 2). Thus, top predators in marine systems tendto have a large body size, low abundance and greaterlongevity than species lower in the food web. Largebodiedtaxa are particularly vulnerable to habitat fragmentation/destruction,but may be less susceptible tocontaminant stress, which hits smaller species disproportionatelyhard. In contrast, there are usually morespecies at lower trophic levels, resulting in greaterfunctional redundancy and more insurance against theeffects of biodiversity loss. These species also havehigher generation rates and, thus, a greater capacityfor adaptive change. Biodiversity loss is, therefore,likely to be distinctly non-random and dependent onbody size, longevity, life-areavulnerability to habitat fragmentationherbivoretop carnivorecarnivoreprimary producervulnerability to stressability to adapt (evolve)species richness (insurance)Fig. 2. Variation in traits, species richness and vulnerabilityto extinction in a model food webthe loss scenario. There is empirical evidence for thedistribution of the traits shown in Fig. 2 (Petchey et al.2004) in freshwater food webs, and an example is providedin Fig. 3 for an estuarine food web.toppredators2avian piscivores>>kg8>kgavianbenthivores1719predatoryfishkggbenthicinvertebrates3505epibenthic predators2avianherbivoresmgc.500307organic matter/microbes phytoplankton macroalgaeFig. 3. Variation in body size (circles)and species richness (numbers) withtrophic position in the Ythan estuaryfood web. From Hall & Raffaelli (1991)and Leaper & Raffaelli (1999)