Fig. 16.

1 Schematic diagram of the Benguela food web adapted from Yodzis

(1998). Many of the linkages and some of the species have been removed for
clarity. What is clear is that there are many linkages between the different
trophic levels in the system and cannibalism (indicated by the circular
arrows) occurs often.
16.2.4 Why systems respond differently to predator removal
The removal of fish predators in marine ecosystems often has limited
consequences for prey species, whereas in many freshwater ecosystems
there are much closer links between predator and prey dynamics (Persson,
Chapter 15, Volume 1). We attribute this to general differences in the
structure of many marine and freshwater ecosystems and suggest that the
indirect effects of fish predator removal in marine systems are smaller and
less predictable than in freshwaters (Jennings and Kaiser 1998). Within lake
ecosystems most biomass is aggregated into a few size classes. These
loosely correspond to phylogenetic groupings such as algae, crustaceans or
fish with a limited range in their biomass spectrum (Sprules et al. 1983;
Sprules and Goyke 1994; Sprules and Stockwell 1995). Conversely, in the
marine environment the biomass spectrum is often extended and there is
more variance in size within the main phylogenetic groupings. Moreover,
phylogenetic groupings tend to contain more species, with a wider range of
life history traits, behaviours and feeding strategies (Jennings and Kaiser
1998).
16.2.5 The collective effects of fishing
Designing experiments to study fishing effects has always been
difficult because it is never clear whether areas subject to different fishing
pressure can be treated as replicates that respond more or less
independently to fishing. Spatial designs have often been used to investigate
the effects of fishing in the tropics, with reefs or islands treated as replicates.

This approach was adopted by Samoilys (1988) and Jennings and Polunin
(1996) to examine the effects of fishing on the structure of fish communities,
and by McClanahan to look at the fish, algal and urchin interactions
described previously. The assumption that different reefs could be treated as
semi-independent replicates was thought to be reasonable because small
(km2) areas protected from fishing, such as marine reserves, would harbour
a diverse and abundant fish community when adjacent reefs were heavily
fished. In addition, fish communities on reefs that were separated by a few
km would respond differently to local fishing pressure. It was also assumed
that the migrations or movements of adult reef-associated fishes between
replicate areas were limited, and this was later confirmed with tagging
studies (e.g. Holland et al. 1993; Zeller and Russ 1998). However, the major
weakness in understanding the level of connectivity between reefs was
associated with the planktonic egg and larval stages (Jones et al., Chapter
16, Volume 1; Polunin, Chapter 14, this volume). These could return to their
natal reef, making the effects of fishing localized, or be transported to reefs
that were many tens to hundreds of km away, making the effects of fishing
collective. Clearly, if the majority of larvae recuited to
sites away from the natal reef, then spatial studies of fishing effects gave
little insight into the longerterm impacts of fishing. Moreover, if we do not
know the scale on which we should test for fishing effects, it is equally
difficult to know the scale on which to impose management strategies. Two
recent studies made direct assessments of larval dispersal rates. Both
concluded that rates of dispersal between reefs were low and that most
larvae were retained close to the natal reef. Swearer et al. (1999) looked at
the evidence for larval retention of bluehead wrasse (Thalassoma
bifasciatum) at St Croix (US Virgin Islands) in the Caribbean. They used
isotopic signatures in the otoliths to identify the source of larvae that were
recruiting to the reefs, and showed that many had been retained close to the
area where they were spawned. Jones et al. (1999) marked the otoliths of
approximately 10 million developing Ambon damselfish (Pomacentrus
amboinensis) eggs with tetracycline, a marker that fluoresces when the
otolith is examined under ultraviolet light. All the eggs were marked on a reef
at Lizard Island on the Great Barrier Reef, and they comprised 0.5 2.5% of
total Ambon damselfish egg production on this reef. Later in the year,
juveniles recruiting to the reef were caught, and their otoliths were removed
and viewed under ultraviolet light. The frequency of otoliths with marks
suggested that 1560% of juveniles returned to the natal reef and that longdistance dispersal was not usual. Thus there is some evidence to suggest
that fish populations are largely self-sustaining on reefs that are separated
by >10km. This suggests that spatial designs can be useful for looking at
longterm fishing effects and that even small and isolated marine reserves
may be useful conservation tools for some reef fish species. (See also
Polunin, Chapter 14, this volume, for further commentary on the degree to
which fish move away from small areas.)

16.3 EFFECTS OF PREY REMOVAL ON SEA BIRDS

Piscivorous sea birds feed primarily on small shoaling fish. In the North
Atlantic, these prey species include herring (Clupea harengus), sprat
(Sprattus sprattus) and sandeels (Ammodytes spp.). Sandeels are
particularly important prey during the breeding and chick rearing season,
whereas clupeoid and gadoid fish become more important dietary
components during the winter period. The International Council for the
Exploration of the Sea (ICES) working group on sea bird ecology (Anon. 1994)
estimated that the annual consumption of fish by 18 sea bird species, based
on sea bird

Fig. 16.2 Estimated total annual consumption of fish and discarded materials
by 18 sea bird species in the North Sea. (Source: data from Anon. 1994.)
abundance estimates and their energetic requirements was 600000
tonnes y-1 (Fig. 16.2). Of this figure the total consumption of small fish,
consisting of sandeel, sprat, young herring and small gadoids, by sea birds is
approximately 250000 tonnes y-1, which is about one-tenth of the small fish
consumed by larger predatory fish (Anon.
1994). It is thought that the abundance of immature fish in the North Sea has
increased due to the overfishing of larger size classes of piscivorous fish (e.g.
Furness and Monaghan 1987; Daan et al. 1990). Similarly, congruent shifts in
sandeel abundance in the western and eastern North Atlantic ecosystems
were also explained by predator removal by fisheries (Sherman et al. 1981).
Although the increase in the amount of small fish intuitively must have
resulted in a larger potential resource for piscivorous sea birds, the
availability of prey may not have changed quite that dramatically. Shifts in
the localized availability of small fish may negatively
affect sea birds even when the wider fish stock is relatively unaffected. For
example, the reproductive output of puffins (Fratercula arctica) that nest on
the Lofoten islands off Norway was reduced over a 20-year period following

the collapse of local herring stocks, even though, elsewhere, herring stocks
remained at relatively high levels (Anker- Nilssen 1992).
16.3.1 Effects of industrial fisheries on sea bird populations
Industrial fisheries in the North Sea have increased substantially since
the early 1960s. These fisheries target species such as sandeels, which are
then converted into animal feed, fertilizer or are even used as fuel. This
fishery has caused concern among conservationists because many top
predators such as seals, cetaceans and sea birds, rely on sandeels as the
main part of their diet (Furness 1990). The direct impacts of fisheries impacts
on prey availability can easily be overlooked as sea birds are able to
compensate for food shortages
by working harder, allocating more time to foraging, switching to alternative
prey types, or
abandoning nesting attempts. Ultimately the impacts of significant
reductions in prey populations become apparent as sea birds starve to death.
During the 1980s, the breeding success of several sea bird species in the
Shetland Islands declined markedly, coincident with a marked decline in
landings of sandeels from an industrial fishery that operated close to the
Shetland coast (Monaghan et al. 1992). The sea bird species affected most
severely were surface feeders such as Arctic terns (Sterna paradisaea),
black-legged kittiwakes (Rissa tridactyla) and great skuas (Catharacta skua)
that fed predominantly on O-group sandeels.