Designing Fish-Tagging Programmes to Understand Fish Movements and Population Dynamics
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Designing fish tagging programmes. Metcalfe et al. NOT TO BE CITED WITHOUT PRIOR REFERENCE TO THE AUTHORS International Council for Use of data storage tags to reveal aspects Exploration of the Seas of fish behaviour CM2006/Q:03 Designing fish-tagging programmes to understand fish movements and population dynamics Julian Metcalfe, D.A. Righton and E. Hunter Centre for Environment, Fisheries and Aquaculture Science, Lowestoft Laboratory, Lowestoft, Suffolk, England UK. tel: + 44 1502 524352, fax: + 44 1502 5513865, e-mail: [email protected] Large-scale fish tagging programmes are becoming more popular as fishery managers realise the importance of including spatial structure in assessment and management models. Two recent EU-funded projects on plaice and cod have shown how information from electronic tags can be used to gain new insights and add value to historic tagging data. Highlights have been the demonstration of unexpected population sub-structuring in plaice, and the realisation that cod behaviour is very variable in response to regional environments. Success does not come without planning and management; of staff, of data and of expectations. We share our experiences from the last 10 years of electronic tagging, and review those of others, to provide an up-to- date analysis of what makes a good tagging programme, and how to get the most from it. Keywords: Data storage tags, migration, behaviour, tagging Designing fish tagging programmes. Metcalfe et al. INTRODUCTION From a management perspective, a fish stock may be defined as that part of a population within which the effects of exploitation on population structure are recognisable (Pawson & Jennings, 1996). If possible, these management (and assessment) stocks should also have a high degree of biological integrity. The biological identification of stocks, and knowledge of their spatial distribution and dynamics, are therefore important aspects of any fisheries management strategy. Many commercially exploited fish species like cod and plaice in European waters are migratory (Metcalfe et al. 2002), making regular seasonal movements between, for example, winter spawning grounds and summer feeding grounds. Theses migrations result in substantial changes in fish stock distribution and so understanding fish migration is clearly important to rational management and conservation of fish stocks. However, obtaining this understanding at appropriate temporal and spatial scales is frequently problematic. Fishermen can provide useful information about seasonal appearances and disappearances of fish, and fisheries statistics can give a general picture of seasonal changes in population distribution. However, such information rarely suffices to describe the migrations of a particular species, and much more detailed information about the behaviour and movements of individuals and populations are needed. But following fish is difficult, particularly in the open sea where, once released, they quickly disappear from view and cannot easily be followed. Mark-recapture studies using various methods of tagging, have been used since the mid 17th century (Walton and Cotton, 1898) as a means of increasing our understanding of fish biology. Mark-recapture methods tell us where individual fish are at two times in their life (i.e. when caught and tagged, and when recaptured). If tagging and recapture are separated by a suitable amount of time (months or even years), it can provide information on stock identity, movements, migration (both rates and routes), abundance, growth, and mortality. Some littoral species, like blennies (family Blenniidae), make seasonal inshore and offshore movements that extend no more than a few kilometres. For such species, large-scale mark-recapture programmes are unlikely to be cost-effective ways of studying their movements. However, species like herring ( Clupea harengus ), mackerel ( Scomber scombrus ), cod (Gadus morhua ), plaice ( Pleuronectes platessa ), Atlantic salmon (genus Salmo), Pacific salmon (genus Oncorhynchus) and eels (Anguilla species), various species of tuna, billfishes and large sharks make more extensive movements over several hundreds or thousands of kilometres. For these species, mark-recapture studies can provide very valuable information about population structure and dynamics. There are many methods for marking or tagging fish (see Parker et al. 1990; Jennings et al. 2001). Branding or fin clipping is a quick and simple way to mark large numbers of fish, and chemical tags such as the tetracycline (an antibiotic which is deposited specifically in calcified areas and fluoresces under ultraviolet light), can easily be applied to large numbers of fish and Designing fish tagging programmes. Metcalfe et al. remain as a permanent mark. Alternatively, various types of tag can be attached to, or placed in, the fish. External tags include Petersen discs used to tag plaice and other flatfishes, while internal tags include the tiny coded wires that are used for the mass marking of young salmon. Simple mark-recapture studies can be very useful for describing gross patterns of population movement, but the method tells us very little about how fish migrate. Population movements derived from mark-recapture studies rely on commercial fishermen reporting details of the time and location of recapture of tagged fish and the results of such studies are therefore inevitably an integration of both fish behaviour and fishing activity that confounds any analysis of population movements. Tagging data can be adjusted for spatial variations in fishing effort, where this is known, but movements of fish into un-fished or un-fishable areas, or changes in fish behaviour which alter availability or catchability, cannot easily be accounted for. It is only by understanding the movements and behaviour of individuals over short (hours and days) medium (days and weeks) and long (seasons and years) time scales, that we can reveal the mechanisms fish use to move about. Understanding these mechanisms allows us to be predictive, rather than simply descriptive. Large-scale fish mark-recapture programmes are therefore becoming more popular as fishery managers realise the importance of including spatial structure in assessment and management models. Two recent EU-funded projects on plaice (EU FAIR Programme, PL96-2079) and cod (CODYSSEY QLRT – 2001 – 00813) have shown how information from electronic tags can be integrated with large mark-recapture tagging programmes to gain new insights and add value to historic tagging data. Highlights have been the demonstration of unexpected population sub- structuring in plaice, and the realisation that cod behaviour is very variable in response to regional environments. Success does not come without planning and management; of staff, of data and of expectations. We share our experiences from the last 10 years of electronic tagging, and review those of others, to provide an up-to-date analysis of what makes a good tagging programme, and how to get the most from it. METHODS Planning a tagging programme Simply wanting to know where fish move to is not, in itself, a good enough reason to embark upon a large mark-recapture programme. Tailoring a tagging programme too closely to existing management regimes and assumptions, however, would be a missed opportunity to collect other useful biological and ecological information. The objectives of a tagging project should therefore be a careful compromise between curiosity-driven research and data collection that will deliver information relevant to fisheries management. Tagging programmes are also multidisciplinary activities, and researchers engaged in them need to be adept in a number of different fields, from publicity to data management. Designing fish tagging programmes. Metcalfe et al. The information that can be applied to fisheries management will vary depending on regional legislation and practice, but the core requirement will be knowledge of the areas occupied by the target stock and the seasonal movements of individuals between them. There are many objectives that can be nested within this core requirement depending on the specific nature of regional management measures. For instance the locations of key habitats such as spawning areas and feeding grounds, the migration routes between them, and patterns of vertical distribution. This kind of information may help to define areas that should be closed seasonally, or fishing gears that should be subject to limitations or exemption (e.g. Thornback rays, Hunter et al. 2004a). Investigating the scope for tagging Having identified the objectives of the tagging programme, it is necessary to understand as much as possible about the target stock, in addition to its environment and fishery, before any further plans can be made. Key to the success of any tagging programme is good recovery of the tag (essential for studies using archival tags), the tagged fish and associated recapture data. For example, it is important to have an idea about the size the "stock", the extent of its geographical distribution, and what tag recovery rates are likely to be. These factors are important in estimating how many fish need to be tagged, where and when, in order to achieve a statistically robust result. In marine fisheries, the area of encounter is potentially vast but can be reduced significantly with backup information from catch data or earlier tagging studies. For electronic tagging programmes, pre-tagging surveys with conventional tags should be carried