Cross-scale observations on distribution and behavioural dynamics of ocean feeding Norwegian spring-spawning herring (Clupea harengus L.)

Interpretation of the behavioural dynamics and distribution of herring requires explicit consideration of spatial and temporal scales since no single mechanism can explain the pattern on all scales. High resolution sonar and echosounders were used as our observation tools. The cluster ratio, mean Nearest Neighbour Distance (mean ISD): mean of the average Inter-school Distance (mean ISD) provides a descriptive index linking scales of distribution pattern within and between clusters of schools. It can be used to compare school clustering patterns for surveys made at different places and seasons. Two mesoscale patterns of school clustering were observed, as indicated by the distribution of nearest neighbour distances; one at 0.05–0.3 km revealed by sonar and another at 0.8–2.5 km revealed by echosounder. In general, schools were tightly aggregated, whilst school clusters were patchily distributed. Heightened feeding motivation explains the predominance of relatively small schools and their tendency to be found closer together and more clustered than large schools. Differences in distribution and density of food patches and predators may account to some degree for the observations (1) schools were distributed according to their size; both small and large schools had nearest neighbours of similar size more often than expected, (2) splitting and joining of schools were the most frequently recorded behavioural events. A considerable amount of variation in distribution and dynamics can be accounted for by changes in behaviour associated with diurnal vertical migrations. Some schools apparently did not perform diurnal vertical migrations. We suggest that when profitable, some herring may ‘‘dip in to the fridge’’ (feed in deep, cold water) during the day rather than pursue food on potentially high energy migrations. 1999 International Council for the Exploration of the Sea

[1]  S. Levin THE PROBLEM OF PATTERN AND SCALE IN ECOLOGY , 1992 .

[2]  D. V. Radakov Schooling in the ecology of fish , 1973 .

[3]  Fish School Size and Migration , 1969 .

[4]  K. Foote Fish target strengths for use in echo integrator surveys , 1987 .

[5]  John D. Neilson,et al.  Diel vertical migrations of marine fishes: an obligate or facultative process? , 1990 .

[6]  M. Keenleyside,et al.  Some Aspects of the Schooling Behaviour of Fish , 1955 .

[7]  E. Simmonds,et al.  Spatial analysis and mapping of acoustic survey data in the presence of high local variability : geostatistical application to North Sea herring (Clupea harengus) , 1996 .

[8]  H. Knudsen The Bergen Echo Integrator: an introduction , 1990 .

[9]  G Sugihara,et al.  Applications of fractals in ecology. , 1990, Trends in ecology & evolution.

[10]  S. Mackinson,et al.  Variation in structure and distribution of pre‐spawning Pacific herring shoals in two regions of British Columbia , 1999 .

[11]  Manfred Milinski,et al.  Predation risk and feeding behaviour , 1993 .

[12]  I. Christensen,et al.  Seasonal distribution, exploitation and present abundance of stocks of large baleen whales (Mysticeti) and sperm whales (Physeter macrocephalus) in Norwegian and adjacent waters , 1992 .

[13]  J. Piatt,et al.  Scale-dependent correlation of seabirds with schooling fish in a coastal ecosystem , 1986 .

[14]  T. Pitcher,et al.  Fish school density and volume , 1979 .

[15]  Ole Arve Misund,et al.  Adaptive behaviour of herring schools in the Norwegian Sea as revealed by high-resolution sonar , 1996 .

[16]  G. Winters Migrations and Activity Levels of Overwintering Atlantic Herring (Clupea harengus harengus) along Southwest Newfoundland , 1977 .

[17]  J. Blaxter,et al.  The Herring: A Successful Species? , 1985 .

[18]  K. Bailey Interaction between the vertical distribution of juvenile walleye pollock Theragra chalcogramma in the eastern Bering Sea, and cannibalism , 1989 .

[19]  M. J. Morgan,et al.  The effect of hunger, shoal size and the presence of a predator on shoal cohesiveness in bluntnose minnows, Pimephales notatus Rafinesque , 1988 .

[20]  A. Magurran,et al.  The adaptive significance of schooling as an anti-predator defense in fish , 1990 .

[21]  I. Røttingen Recent migration routes of Norwegian spring spawning herring , 1992 .

[22]  J. Godin,et al.  Foraging on patchily distributed prey by a cichlid fish (Teleostei, Cichlidae): A test of the ideal free distribution theory , 1984, Animal Behaviour.

[23]  Matthew T. Wilson,et al.  Mesoscale acoustic patterns of juvenile walleye pollock (Theragra chalcogramma) in the western Gulf of Alaska , 1996 .

[24]  I. Christensen,et al.  Occurrence and diet of killer whales in northern Norway: seasonal patterns relative to the distribution and abundance of Norwegian spring-spawning herring , 1996 .

[25]  U. Lindstrøm,et al.  Diet and food availability for Northeast Atlantic minke whales Balaenoptera acutorostrata , 1995 .

[26]  J. Krebs,et al.  Arms races between and within species , 1979, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[27]  Magnar Aksland,et al.  Schooling dynamics of norwegian spring spawning herring (Clupea harengus L.) in a coastal spawning area , 1996 .

[28]  Michael W. Davis,et al.  Behavioral responses of juvenile walleye pollock Theragra chalcogramma Pallas to light, thermoclines and food : possible role in vertical distribution , 1990 .

[29]  Øyvind Ulltang,et al.  Varlation in migration and abundance of norwegian spring spawning herring (Clupea harengus L.) , 1997 .

[30]  O L E A Rv,et al.  Underwater acoustics in marine fisheries and fisheries research , 1997 .

[31]  T. J. Pitcher,et al.  Hunger motivation as a promoter of different behaviours within a shoal of herring: selection for homogeneity in fish shoal? , 1989 .

[32]  Ole Arve Misund,et al.  THE CHALLENGE OF THE HERRING IN THE NORWEGIAN SEA : MAKING OPTIMAL COLLECTIVE SPATIAL DECISIONS , 1998 .

[33]  Rune Vabø,et al.  An individual based model of fish school reactions: predicting antipredator behaviour as observed in nature , 1997 .

[34]  F. Arenas,et al.  Diel vertical and offshore-inshore movements of anchovies off the central Baja California coast , 1995 .

[35]  S. L. Lima,et al.  Behavioral decisions made under the risk of predation: a review and prospectus , 1990 .

[36]  Carlos J. Robinson,et al.  Food competition in a shoal of herring: The role of hunger , 1994 .

[37]  Susan M. Sogard,et al.  Effects of light, thermoclines and predator presence on vertical distribution and behavioral interactions of juvenile walleye pollock, Theragra chalcogramma Pallas , 1993 .

[38]  Ingolf Røttingen,et al.  Feeding and migration of Norwegian spring spawning herring in the Norwegian Sea , 1994 .

[39]  Ole Arve Misund,et al.  Tracking herring schools with a high resolution sonar. Variations in horizontal area and relative echo intensity , 1998 .

[40]  T. Pitcher,et al.  Shoal size, patch profitability and information exchange in foraging goldfish , 1983, Animal Behaviour.

[41]  Ole Arve Misund,et al.  Mapping the shape, size, and density of fish schools by echo integration and a high-resolution sonar , 1995 .

[42]  Ole Arve Misund,et al.  Dynamics of moving masses: variability in packing density, shape, and size among herring, sprat, and saithe schools , 1993 .

[43]  J. Blaxter,et al.  The Biology of the Clupeoid Fishes , 1982 .

[44]  D. Skagen,et al.  Improved mapping of schooling fish near the surface: comparison of abundance estimates obtained by sonar and echo integration , 1996 .

[45]  Aril Slotte Relations between seasonal migrations and fat content in Norwegian spring spawning herring (Clupea harengus L.) , 1996 .

[46]  Ole Arve Misund,et al.  Trawl designs and techniques used by Norwegian research vessels to sample fish in the pelagic zone , 1995 .

[47]  A. J. Paul,et al.  Effect of food intake and temperature on growth and conversion efficiency of juvenile walleye pollock (Theragra chalcogramma (Pallas)): a laboratory study , 1986 .