The organising principles behind adaptive collective formations in animals are not well understood. Here we document that persistent vertical hourglass formations occur during the critical spawning period in huge shoals of the herring Clupea harengus spawning demersally. We compared individual maturity stage between the upper and lower components of the formations and found the proportion of spawners to pre-and post-spawners to be significantly higher in the lower than in the upper components. Our results provide empirical support for the hypothesis that these formations are a collective output of individual risk assessment in a compromise between survival and reproduction, where non-spawners minimise the time spent in the demersal zone, where density of gadoid predators was found to be highest. Conflicting individual motivation may thus be an organising principle behind mass formations in fish shoals.
KEY WORDS: Herring · Spawning · Hourglass formation · MotivationResale or republication not permitted without written consent of the publisher Mar Ecol Prog Ser 358: 295-300, 2008 spawners to be lowest in the low component of the hourglasses and the density of predators to be highest near the bottom.
MATERIALS AND METHODSWe analysed time series of acoustic data from the Norwegian spring spawning (NSS) herring Clupea harengus spawning areas along the Norwegian coast, where each spring herring are found in some of the largest known animal aggregations on the globe. The acoustic data material was acquired in February and March 1994 to 2006 (except for the years 1996 in part as abundance estimate surveys of NSS herring, and in part as distribution mapping and/or behavioural studies within selected herring spawning sites (Fig. 1). In total, 260 trawl stations were available, and acoustic data were collected during 170 d at sea for an area covering roughly 30 000 nautical miles (n miles).The collection of data was carried out by experienced personal and followed a standard scheme: Acoustic recording was done with a 38 kHz SIMRAD EK 500 echo sounder and integrator (replaced with EK 60 in 2005) and post-processed using the Bergen Echo Integrator (BEI) (Foote et al. 1991). Acoustic signal characteristics combined with trawl catches were used for reference when allocating acoustic densities to species. All biological samples were obtained by pelagic trawl upon recordings of herring and not at random or fixed stations. If permitted by the size of the catch, 100 herring were randomly sampled from each trawl haul. Herring with maturing and mature gonads were categorised as 'pre-spawners', individuals with running gonads as 'spawners' and individuals with empty or quasi-empty gonads as 'post-spawners'. The acoustic density values were stored by species in nautical area scattering coefficient (NASC) units (MacLennan et al. 2002) in a database with a horizontal resolution of 1 n mile and a vertical resolution of 10 m, referenced to the surface.In order to distinguish potential hourglass formations, we first searched...