The Functioning of Marine Ecosystems A fisheries perspective

1. The Functioning of Marine EcosystemsA fisheries perspective Philippe Cury Lynne Shannon Yunne-Jai Shin

2. Definition of Marine Ecosystems • The term “Ecosystem” is recent (Tansley 1935) • An ecosystem is defined as “a spatially explicit unit of earth that includes all of the organisms, along with all the components of the abiotic environment within its boundaries” (Likens 1992) • A marine ecosystem contains water, detritus, hundreds of kinds of organisms including bacteria, phytoplankton, zooplankton, fishes, mammals, birds … and fishers!

3. Ecosystem as an ‘Uncomfortable’ large scale Unit for Ecological studies but an Integrative level for Fisheries Management • Ecosystems have no apparent boundaries and lack the sort of clear objective or purpose that can be ascribed to other, more tractable, biological or ecological entities (e.g. cell, individual or population): there is no clear objective function to maximize • However the ecosystem is now viewed as an integrative level, and its overall complexity is perceived as critical to its sustainability

4. Questions Regarding the Exploitation of Marine Ecosystems : • Which species are most critical, and which ecological processes are most sensitive to exploitation? • Does the removal of top predators have a strong impact on lower trophic levels? • Does heavy exploitation of forage species, such as anchovies and sardines, cause changes in the functioning of upwelling ecosystems?

5. The key to answering these questions and exploring whether general principles apply lies in understanding the energy flow in the ecosystem

6. Which Energy Flows in Marine Ecosystems? Who is controlling Whom? Top- Down Bottom-Up Wasp-Waist

7. Bottom-up Control

8. The Very Small drive the Very Large

9. westerly weather Parallel long-term trends across four marine trophic levels and weather in the North Sea(from Aebischer et al, Nature 1990) phytoplankton zooplankton herring Kittiwake laydate Kittiwake clutch Kittiwake chicks

10. Five ecosystems in the Pacific were affected by the mid-1970s climate change event (from McGowan et al. 1998)

11. Regime Shifts within the Peruvian Ecosystem(from Pauly and Tsukayama 1987)

12. Bottom-up Control • Under bottom-up control, the physical environment drastically affects the overall productivity (i.e. the carrying capacity) of ecosystems, but more importantly the dynamics of fish assemblages in a more or less predictable way… • Decadal-scale regime shifts suggest the existence of multiple stable states in fish communities, resulting in sustained or un-sustained fisheries and induced ecosystem changes • A Bottom-up control offers a comprehensive framework for understanding how different components would react to environmental changes or to changes at the bottom of the food chain

13. Top-Down Control

14. The Very Large drive the Very Small

15. Bigger Fish Eat Smaller Fish

16. Predators are constrained by the size of their jaw and prey fish that are less than 1/3 their own size Who is eating whom? Cannibalism, Omnivory The relative stability of the total fish biomass compared to that of individual species Size-based predation provides an explanation for observed size spectra in marine ecosystems

17. Trophic Cascade in Alaska resulting in inverse patterns in abundance or biomass across trophic links in a food web(from Science, Estes et al. 1998)

18. Top-Down Control • A top-down control can help to understand several observed ecological patterns at an ecosystem level when removing top predators • Not all cascades propagate to lower trophic levels or have significant impacts on ecosystem processes as numerous compensatory mechanisms dampen or eliminate them • Fishing usually greatly reduces the abundance of top predators, and it stands to reason that the abundance of prey populations and their effects on marine communities will increase after release from predator control

19. Wasp-Waist Control

20. Small pelagic fishes drive both the very large and the very small

21. Birds and Sardine in South Africa and Namibia(from Crawford 1999)

22. Black-Sea Pelagic fish abundance & zooplankton 300 Zooplanktonabundance 200 100 0 0 200 400 600 800 Ghana Pelagic fish abundance 200 South-Africa 100 7 0 100 200 300 5 0 Japan 3 60 100 300 500 700 40 20 0 2000 4000 6000 0

23. Wasp-Waist Control • Under wasp-waist control the collapse of a dominant prey species can generate drastic changes at higher, but most surprisingly at lower trophic levels • As fisheries remove substantial amounts of small pelagic fish one must carefully consider the implications for the other species in the ecosystem

24. Bottom-Up Top- Down Wasp-Waist

25. Who is controlling whom in Marine Food-Webs ?

26. No General Theory can be ascribed to the Functioning of Marine Ecosystems the ability to predict ecosystem behavior is limited

27. However ecosystems are neither totally predictable nor totally unpredictable : tentative generalisations can be proposed • Control by the environment (bottom-up control) predominates • Control by predators (top-down control) plays a role in dampening ecosystem-level fluctuations • Trophic cascades are seldom found, except in lakes, or in marine hard substrata ecosystems and mainly for less complex food-webs • Wasp-waist control is most probable in upwelling systems.

28. From ‘common sense’ and ‘pet concept’ toward an operational framework for dealing with responsible fisheries in marine ecosystems? • These difficulties do not mean that an ecosystem approach to fisheries management should be abandoned or that we should just wait for more additional results on the functioning of ecosystems • Major steps are urgently needed, that will define an operational framework for dealing with responsible fisheries in marine ecosystems, for example by using ecosystem-based indicators and reference points • This is a complex issue that needs to integrate our simplistic and disparate views of nature and to reach major steps towards incorporating our recent and incomplete, but consequential, theoretical background on the functioning of marine ecosystems

29. ‘Not only is the science incomplete, but the [eco]system itself is a moving target, evolving because of the impact of management and the progressive expansion of the scale of human influences on the planet’Holling C.S. (1995)