Strategies of space occupation by anchovy and sardine in the southern Benguela: the role of stock size and intra-species competition

Strategies of space occupation by anchovy and sardine populations in the southern Benguela during cruises in 1994 (low biomass) and 2001 (high biomass) are investigated using geostatistical tools. The spatial expansion of anchovy and sardine during the high-density year resulted in an increase in the number and density of hotspots and, in the case of anchovy, a relative increase in the contribution of mid- to high-density intervals to the overall survey biomass. Variogram autocorrelation ranges were larger in the high-biomass year. Otherwise, similarities and differences in the strategies of space occupation between anchovy and sardine are consistent between years of low and high biomass, and are therefore density-independent. Variograms of indicator variables revealed comparable structures for both species and years, with autocorrelation ranges of 30 nautical miles for the smallest indicator (0.1 g m(-2)) and 10 miles for the largest (100 g m(-2)), reflecting intrinsic spatial structures independent of stock size. The ideal survey inter-transect distance, defined as the autocorrelation range of the highest density indicator, was not influenced by the expanded distribution of both species. Cumulative frequency distributions and cross-variograms revealed differences in space occupancy between both species. Despite the areal expansion, and in contrast to anchovy, a significant portion of the sardine biomass continued to be contributed by a few very high-density areas with little spatial structure, as indicated by large nugget effects in the variograms and lack of transitions in the cross-variograms. Both species were spatially segregated in the low-biomass year, but overlapped extensively in the high-biomass year. (c) 2004 International Council for the Exploration ofthe Sea. Published by Elsevier Ltd. All rights reserved.