Toward a Solution of the “Peruvian Puzzle”: Pelagic Food-Web Structure and Trophic Interactions in the Northern Humboldt Current Upwelling System Off Peru

The northern Humboldt Current upwelling system (HCS) belongs to the most productive marine ecosystems, providing five to eight times higher fisheries landings per unit area than other coastal upwelling systems. To solve this “Peruvian puzzle”, to elucidate the pelagic food-web structure and to better understand trophic interactions in the HCS, a combined stable isotope and fatty acid trophic biomarker approach was adopted for key zooplankton taxa and higher trophic positions with an extensive spatial coverage from 8.5 to 16°S and a vertical range down to 1,000 m depth. A pronounced regional shift by up to ∼5‰ in the δ15N baseline of the food web occurred from North to South. Besides regional shifts, δ15N ratios of particulate organic matter (POM) also tended to increase with depth, with differences of up to 3.8‰ between surface waters and the oxygen minimum zone. In consequence, suspension-feeding zooplankton permanently residing at depth had up to ∼6‰ higher δ15N signals than surface-living species or diel vertical migrants. The comprehensive data set covered over 20 zooplankton taxa and indicated that three crustacean species usually are key in the zooplankton community, i.e., the copepods Calanus chilensis at the surface and Eucalanus inermis in the pronounced OMZ and the krill Euphausia mucronata, resulting in an overall low number of major trophic pathways toward anchovies. In addition, the semi-pelagic squat lobster Pleuroncodes monodon appears to play a key role in the benthic-pelagic coupling, as indicated by highest δ13C’ ratios of −14.7‰. If feeding on benthic resources and by diel vertical migration, they provide a unique pathway for returning carbon and energy from the seafloor to the epipelagic layer, increasing the food supply for pelagic fish. Overall, these mechanisms result in a very efficient food chain, channeling energy toward higher trophic positions and partially explaining the “Peruvian puzzle” of enormous fish production in the HCS.

Flooding affects food web structure and basal sources supporting fish guilds in a subtropical wetland and shallow lake

Aquatic ecosystems exchange nutrients and organic matter with surrounding terrestrial ecosystems, and floods import allochthonous material from riparian areas into fluvial systems. We surveyed food web components of a wetland and shallow lake in a subtropical coastal region of Brazil to examine how community trophic structure and the entrance of allochthonous material into the food web were affected by floods. Stable isotope analysis was performed for samples of terrestrial and aquatic basal production sources and aquatic animals to trace the origin of organic matter assimilated by aquatic animals and estimate vertical trophic positions and food chain length. Lake and wetland trophic structures were compared for cool/wet and warm/dry seasons. Food web structure was hypothesized to differ based on hydrology, with the more stable lake having greater food web complexity, and seasonal flooding resulting in greater allochthonous inputs to the aquatic food web. We compared spatial and temporal variation in assemblage trophic structure using an adapted isotopic ellipse approach that plots assemblage elements according to δ13C on the x-axis and estimated TP on the y-axis. Lake trophic structure was more complex with longer food chains compared to that of the wetland. A greater contribution from terrestrial resources to animal biomass was observed in the wetland during the cool/wet period, and food chains in both habitats tended to be longer during the cool/wet period. Findings supported the hypothesis of greater assimilation of allochthonous sources during floods and greater trophic complexity in the more hydrologically stable system.

Less is worse than none: ineffective adaptive foraging can destabilise food webs

Consumers can potentially adjust their diet in response to changing resource abundances, thereby achieving better foraging payoffs. Although previous work has explored how such adaptive foraging scales up to determine the structure and dynamics of food webs, consumers may not be able to perform perfect diet adjustment due to sensory or cognitive limitations. Whether the effectiveness of consumers' diet adjustment alters food-web consequences remains unclear. Here, we study how adaptive foraging, specifically the effectiveness (i.e. rate) with which consumers adjust their diet, influences the structure, dynamics, and overall species persistence in synthetic food webs. We model metabolically-constrained optimal foraging as the mechanistic basis of adaptive diet adjustment and ensuing population dynamics within food webs. We compare food-web dynamical outcomes among simulations sharing initial states but differing in the effectiveness of diet adjustment. We show that adaptive diet adjustment generally makes food-web structure resilient to species loss. Effective diet adjustment that maintains optimal foraging in the face of changing resource abundances facilitates species persistence in the community, particularly reducing the extinction of top consumers. However, a greater proportion of intermediate consumers goes extinct as optimal foraging becomes less-effective and, unexpectedly, slow diet adjustment leads to higher extinction rates than no diet adjustment at all. Therefore, food-web responses cannot be predicted from species' responses in isolation, as even less-effective adaptive foraging benefits individual species (better than non-adaptive) but can harm species' persistence in the food web as a whole (worse than non-adaptive). Whether adaptive foraging helps or harms species coexistence has been contradictory in literature. Our finding that it can stabilise or destabilise the food web depending on how effectively it is performed help reconcile this conflict. Inspired by our simulations, we deduce that there may exist a positive association between consumers' body size and adaptive-foraging effectiveness in the real world. We also infer that such effectiveness may be higher when consumers cognise complete information about their resources, or when trophic interactions are driven more by general traits than by specific trait-matching. We thereby suggest testable hypotheses on species persistence and food-web structure for future research, in both theoretical and empirical systems.