Ecological and acoustic investigations of jellyfish (Scyphozoa and Hydrozoa)
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As the biomass of jellyfish (medusae of the Scyphozoa and Hydrozoa) has risen in numerous locations worldwide, awareness of their potential to exert a controlling influence on marine ecosystems and hinder the recruitment of fish stocks has increased. Medusae are capable of intensive, size–selective, predation on zooplankton, which may alter the composition of the plankton community. Jellyfish are often found in dense layers, up to hundreds of metres thick, which can extend horizontally for hundreds of kilometres. Such aggregations may benefit specialist feeders, such as turtles, that rely upon jellyfish for food and those fish that are able to find refuge under the jellyfish umbrellas. Nonetheless, the predominance of jellyfish in pelagic ecosystems is not generally viewed as desirable; in fact, this situation has been portrayed as the result of pollution and overexploitation of otherwise productive seas. However, jellyfish are sensitive indicators of environmental change, and their populations appear to respond to climatic fluctuations, so jellyfish warrant study for their intrinsic ecosystem role particularly given present concerns over climate change. With growing acceptance that fishery management should take an holistic ‘ecosystem approach’, knowledge of the interactions between jellyfish, fisheries and climate may be vital in progression towards the goal of ecosystem–based sustainable management of fisheries. Unfortunately, due to their gelatinous nature, medusae are difficult to sample using conventional netting techniques and data on changes in distribution and abundance are consequently sparse. Recent studies have demonstrated that medusae can be detected acoustically and that this technique could provide a rapid and cost–effective estimate of their biomass and distribution. This thesis reports my endeavour to demonstrate the ecosystem role of medusae and to develop acoustic techniques to monitor their biomass. Through regession analyses, I link the abundance of medusae (Aurelia aurita, Cyanea lamarckii, and Cyanea capillata) in regions of the North Sea to climatic fluctuations, as quantified by the North Atlantic Oscillation Index, and show that medusae may be important indicators of regional ecosystem change. The mechanisms linking climatic fluctuations to ecosystem changes are explored via a correlative modelling approach using General Additive Models; I show that the mechanisms are location dependent and explainable in terms of direct, rapidly responding (intra–annual) influences (surface warming, river run–off, and wind–driven mixing and advection) and longer–term (interannual) oceanographic responses (changes in circulation currents i.e. the northward extent of the gulf stream and relative strength of inflow into the North Sea of the North Atlantic current, Continental Shelf Jet and Arctic waters). I present correlative evidence for a detrimental impact by Aurelia aurita on herring 0–group recruitment, once the influence of interannual change in herring spewing stock biomass on recruitment is factored out through modelling with a Ricker stock–recruitment relationship. Similarly, a commensal relationship between whiting and Cyanea spp. medusae is shown to improve North Sea whiting survival to the 1–group. In progress towards the automated acoustic identification of species, I have developed an in situ discrimination tool that can distinguish between echoes from: Aequorea aequorea; Chrysaora hysoscella; clupeid fish (sardine, anchovy and round herring); and horse mackerel/Cape hake. The technique relies upon characteristic differences in echo strength between frequencies, which are determined for each jellyfish species and finfish group using combined multifrequency acoustic and pelagic trawl samples. This method has facilitated the world’s first acoustic–based estimate of jellyfish biomass in the Namibian Benguela Sea. The 12.2 million tonnes of biomass of medusae (Aequorea aequorea and Chrysaora hysoscella) in the Namibian Benguela Sea was found to be greater than the combined biomass, 3.6 million tonnes, of commercially important fish (horse mackerel, Cape hake, sardines, anchovy, and round herring) in the same area. These results suggest that medusae may have an important role in the Benguela ecosystem that has previously been overlooked and that their biomass should be monitored.
Thesis, PhD Doctor of Philosophy
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