Trophic ecology of jumbo squid and predatory fishes in the Northern Humboldt Current System

This work provides a contribution to a better understanding of the trophic ecology of important predators in the Northern Humboldt Current System, the jack mackerel (Trachurus murphyi), the chub mackerel (Scomber japonicus) and the jumbo squid (Dosidicus gigas) by the characterization of the highly variable feeding patterns of these species at different spatiotemporal scales. We provided new knowledge on the comparative trophic behaviour of these species, defined as opportunistic in previous investigations. For that purpose we applied a variety of statistical methods to an extensive dataset of 27,188 non-empty stomachs. We defined the spatial organization of the forage fauna of these predators and documented changes in prey composition according to predators’ size and spatiotemporal features of environment. Our results highligh the key role played by the dissolved oxygen. We also deciphered an important paradox on the jumbo squid diet: why do they hardly forage on the huge anchovy (Engraulis ringens) biomass distributed of coastal Peru? We showed that the shallow oxygen minimum zone present off coastal Peru could hamper the co-occurrence of jumbo squids and anchovies. In addition, we proposed a conceptual model on jumbo squid trophic ecology including the ontogenetic cycle, oxygen and prey availability. Moreover we showed that the trophic behaviour of jack mackerel and chub mackerel is adapted to forage on more accessible species such as for example the squat lobster Pleurocondes monodon and Zoea larvae. Besides, both predators present a trophic overlap. But jack mackerel was not as oracious as chub mackerel, contradictorily to what was observed by others authors. Fish diet presented a high spatiotemporal variability, and the shelf break appeared as a strong biogeographical frontier. Diet composition of our fish predators was not necessarily a consistent indicator of changes in prey biomass. El Niño events had a weak effect on the stomach fullness and diet composition of chub mackerel and jack mackerel. Moreover, decadal changes in diet diversity challenged the classic paradigm of positive correlation between species richness and temperature. Finally, the global patterns that we described in this work, illustrated the opportunistic foraging behaviour, life strategies and the high degree of plasticity of these species. Such behaviour allows adaptation to changes in the environment.

End-to-end modelling for an ecosystem approach to fisheries in the Northern Humboldt Current Ecosystem

This work represents an original contribution to the methodology for ecosystem models' development as well as the rst attempt of an end-to-end (E2E) model of the Northern Humboldt Current Ecosystem (NHCE). The main purpose of the developed model is to build a tool for ecosystem-based management and decision making, reason why the credibility of the model is essential, and this can be assessed through confrontation to data. Additionally, the NHCE exhibits a high climatic and oceanographic variability at several scales, the major source of interannual variability being the interruption of the upwelling seasonality by the El Niño Southern Oscillation, which has direct e ects on larval survival and sh recruitment success. Fishing activity can also be highly variable, depending on the abundance and accessibility of the main shery resources. This context brings the two main methodological questions addressed in this thesis, through the development of an end-to-end model coupling the high trophic level model OSMOSE to the hydrodynamics and biogeochemical model ROMS-PISCES: i) how to calibrate ecosystem models using time series data and ii) how to incorporate the impact of the interannual variability of the environment and shing. First, this thesis highlights some issues related to the confrontation of complex ecosystem models to data and proposes a methodology for a sequential multi-phases calibration of ecosystem models. We propose two criteria to classify the parameters of a model: the model dependency and the time variability of the parameters. Then, these criteria along with the availability of approximate initial estimates are used as decision rules to determine which parameters need to be estimated, and their precedence order in the sequential calibration process. Additionally, a new Evolutionary Algorithm designed for the calibration of stochastic models (e.g Individual Based Model) and optimized for maximum likelihood estimation has been developed and applied to the calibration of the OSMOSE model to time series data. The environmental variability is explicit in the model: the ROMS-PISCES model forces the OSMOSE model and drives potential bottom-up e ects up the foodweb through plankton and sh trophic interactions, as well as through changes in the spatial distribution of sh. The latter e ect was taken into account using presence/ absence species distribution models which are traditionally assessed through a confusion matrix and the statistical metrics associated to it. However, when considering the prediction of the habitat against time, the variability in the spatial distribution of the habitat can be summarized and validated using the emerging patterns from the shape of the spatial distributions. We modeled the potential habitat of the main species of the Humboldt Current Ecosystem using several sources of information ( sheries, scienti c surveys and satellite monitoring of vessels) jointly with environmental data from remote sensing and in situ observations, from 1992 to 2008. The potential habitat was predicted over the study period with monthly resolution, and the model was validated using quantitative and qualitative information of the system using a pattern oriented approach. The nal ROMS-PISCES-OSMOSE E2E ecosystem model for the NHCE was calibrated using our evolutionary algorithm and a likelihood approach to t monthly time series data of landings, abundance indices and catch at length distributions from 1992 to 2008. To conclude, some potential applications of the model for shery management are presented and their limitations and perspectives discussed.

Carbon cycling in the northern Humboldt Current off Chile

The structure of the zooplankton foodweb and their dominant carbon fluxes were studied in the upwelling system off northern Chile (Mejillones Bay; 23°S) between October 2000 and December 2002. High primary production (PP) rates (18 gC/m**2 d) were mostly due to the net-phytoplankton size fraction (>23 µm). High PP has been traditionally associated with the wind-driven upwelling fertilizing effect of equatorial subsurface waters, which favour development of a short food chain dominated by a few small clupeiform fish species. The objective of the present work was to study the trophic carbon flow through the first step of this 'classical chain' (from phytoplankton to primary consumers such as copepods and euphausiids) and the carbon flow towards the gelatinous web composed of both filter-feeding and carnivorous zooplankton. To accomplish this objective, feeding experiments with copepods, appendicularians, ctenophores, and chaetognaths were conducted using naturally occurring plankton prey assemblages. Throughout the study, the total carbon ingestion rates showed that the dominant appendicularian species and small copepods consumed an average of 7 and 5 µgC/ind d, respectively. In addition, copepods ingested particles mainly in the size range of nano- and microplankton, whereas appendicularians ingested in the range of pico- and nanoplankton. Small copepods and appendicularians removed a small fraction of total daily PP (range 6-11%). However, when the pico- + nanoplankton fractions were the major contributors to total PP (oligotrophic conditions), grazing by small copepods increased markedly to 86% of total PP. Under these more oligotrophic conditions, the euphausiids grazing increased as well, but only reached values lower than 5% of total PP. During this study, chaetognaths and ctenophores ingested an average of 1 and 14 copepods/ind d, respectively. In terms of biomass consumed, the potential impact of carnivorous gelatinous zooplankton on the small-size copepod community (preferred prey) was important (2-12% of biomass removed daily). However, their impact produced more significant results on copepod abundance (up to 33%), which suggests that carnivorous gelatinous zooplankton may even modulate (control) the abundance of some species as well as the size structure of the copepod community.

Evolution of 1996-1999 La Niña and El Niño Conditions Off the Western Coast of South America: A Remote Sensing Perspective

We present the evolution of oceanographic conditions off the western coast of South America between 1996 and 1999, including the cold periods of 1996 and 1998-1999 and the 1997-1998 El Niño, using satellite observations of sea level, winds, sea surface temperature (SST), and chlorophyll concentration. Following a period of cold SST and low sea levels in 1996, both were anomalously high between March 1997 and May 1998. The anomalies were greatest between 5 degrees S and 15 degrees S, although they extended beyond 40 degrees S. Two distinct peaks in sea level and SST occurred in June-July 1997 and December 1997 to January 1998, separated by a relaxation period (August-November) of weaker anomalies. Satellite winds were upwelling favorable throughout the time period for most of the region and in fact increased between November 1997 and March 1998 between 5 degrees S and 25 degrees S. Satellite-derived chlorophyll concentrations are available for November 1996 to June 1997 (Ocean Color and Temperature Sensor (OCTS)) and then from October 1997 to present (Sea-viewing Wide Field-of-view Sensor (SeaWiFS)). Near-surface chlorophyll concentrations fell from May to June 1997 and from December 1997 to March 1998. The decrease was more pronounced in northern Chile than off the coast of Peru or central Chile and was stronger for larger cross-shelf averaging bins since nearshore concentrations remained relatively high.

Pollen record from sediment core M77/2_056-3 (Bay of Guayaquil, Peru)

We analyzed the pollen content of a marine core located near the bay of Guayaquil in Ecuador to document the link between sea surface temperatures (SST) and changes in rainfall regimes on the adjacent continent during the Holocene. Based on the expansion/regression of five vegetation types, we observe three successive climatic patterns. In the first phase, between 11,700 and 7700 cal yr BP, the presence of a cloud (Andean) forest in the mid altitudes and mangroves in the estuary of the Guayas Basin, were associated with a maximum in boreal summer insolation, a northernmost position of the Intertropical Convergence Zone (ITCZ), a land- sea thermal contrast, and dryness. Between 7700 and 2850 cal yr BP, the expansion of the coastal herbs and the regression of the mangrove indicate a drier climate with weak ITCZ and low ENSO variability while austral winter insolation gradually increased. The interval between 4200 and 2850 cal yr BP was marked by the coolest and driest climatic conditions of the Holocene due to the weak influence of the ITCZ and a strengthening of the Humboldt Current. After 2850 cal yr BP, high variability and amplitude of the Andean forest changes occurred when ENSO frequency and amplitude increased, indicating high variability in land-sea connections. The ITCZ reached the latitude of Guayaquil only after 2500 cal yr BP inducing the bimodal precipitation regime we observe today. Our study shows that besides insolation, the ITCZ position and ENSO frequency, changes in eastern equatorial Pacific SSTs play a major role in determining the composition of the ecosystems and the hydrological cycle of the Ecuadorian Pacific coast and the Western Cordillera in Ecuador.

Epibiosis between the sand crab Emerita analoga and the mussel Semimytilus algosus

Coastal zones of the Humboldt Current Upwelling System (HCUS) are composed both of rocky and sandy beaches inhabited by macrozoobenthic communities. These show oscillating changes in the dominance of species; the abundance of the sand crab Emerita analoga is linked to phases of the El Niño Southern Oscillation (ENSO). The biogenic surfaces of these crabs serve as substrate for opportunistic colonizers. This study is the first record of an epibiosis between E. analoga and the rock mussel Semimytilus algosus, detected at a southern Peruvian sandy beach. Mussels fouled a wide size-range of adult E. analoga (7.3%) but they themselves belonged to small-size classes. The largest S. algosus was 17.4 mm in length. Highest permanence of epibionts was found on larger sand crabs (maximum between 24 and 27 mm). Significantly more mussels were found on the ventral surface (39.4%) compared to 10 other surface areas of the sand crab. Possible benefits and disadvantages of the observed epibiosis for both the basibiont and the epibiont are discussed.

The Southern Ocean and South Pacific Region

The Region comprises three sub-regions (FAO Statistical Areas) with very different characteristics. The South Pacific includes the vast and virtually unpopulated Southern Ocean surrounding the Antarctic. It has the world’s largest fisheries off Peru and Chile and some of the world’s best managed fisheries in Australia and New Zealand. The Region has over 27% of the world’s ocean area and over 98% of the Region’s total area of 91 million km2 is ‘open ocean’. The Region contains less than 5% of the global continental shelf area and only a fraction of this area is covered by three large marine ecosystems (the New Zealand Shelf, the Humboldt Current and the Antarctic large marine ecosystems (LMEs). The Humboldt Current System (HCS) is the world’s largest upwelling which provides nutrients for the world’s largest fisheries. The Region also has a high number of seamounts. The marine capture fisheries of the Region produce over 13 million tons annually and an expanding aquaculture industry produces over 1.5 million tons. Peru’s anchoveta fishery provides about half the world’s supply of fish meal and oil, key ingredients of animal and fish feeds. El Niño Southern Oscillations (ENSOs), known more generally as El Niños, can substantially change the species composition of the key small pelagic catches (anchovy, sardine, horse mackerel and jack mackerel) causing production to fluctuate from about 4-8 million tons. Partly due to the lack of upwelling and shelf areas, fisheries production in the Southern Ocean and Area 81 is relatively small but supports economically important commercial and recreational fisheries and aquaculture in New Zealand and in New South Wales (Australia). Krill remains a major underexploited resource, but is also a keystone species in the Antarctic food web. The Region is home to numerous endangered species of whales, seals and seabirds and has a high number of seamounts, vulnerable ecosystems fished for high-value species such as orange roughy.

The Southern Ocean and South Pacific Region

The Region comprises three sub-regions (FAO Statistical Areas) with very different characteristics. The South Pacific includes the vast and virtually unpopulated Southern Ocean surrounding the Antarctic. It has the world’s largest fisheries off Peru and Chile and some of the world’s best managed fisheries in Australia and New Zealand. The Region has over 27% of the world’s ocean area and over 98% of the Region’s total area of 91 million km2 is ‘open ocean’. The Region contains less than 5% of the global continental shelf area and only a fraction of this area is covered by three large marine ecosystems (the New Zealand Shelf, the Humboldt Current and the Antarctic large marine ecosystems (LMEs). The Humboldt Current System (HCS) is the world’s largest upwelling which provides nutrients for the world’s largest fisheries. The Region also has a high number of seamounts. The marine capture fisheries of the Region produce over 13 million tons annually and an expanding aquaculture industry produces over 1.5 million tons. Peru’s anchoveta fishery provides about half the world’s supply of fish meal and oil, key ingredients of animal and fish feeds. El Niño Southern Oscillations (ENSOs), known more generally as El Niños, can substantially change the species composition of the key small pelagic catches (anchovy, sardine, horse mackerel and jack mackerel) causing production to fluctuate from about 4-8 million tons. Partly due to the lack of upwelling and shelf areas, fisheries production in the Southern Ocean and Area 81 is relatively small but supports economically important commercial and recreational fisheries and aquaculture in New Zealand and in New South Wales (Australia). Krill remains a major underexploited resource, but is also a keystone species in the Antarctic food web. The Region is home to numerous endangered species of whales, seals and seabirds and has a high number of seamounts, vulnerable ecosystems fished for high-value species such as orange roughy.

Hydrographic Conditions Off Northern Chile During the 1996-1998 La Niña and El Niño Events

The evolution of oceanographic conditions in the upwelling region off northern Chile (18 degrees-24 degrees S) between 1996 and 1998 (including the 1997-1998 El Niño) is presented using hydrographic measurements acquired on quarterly cruises of the Chilean Fisheries Institute, with sea surface temperature (SST), sea level, and wind speeds from Arica (18.5 degrees S), Iquique (20.5 degrees S), and Antofagasta (23.5 degrees S) and a time series of vertical temperature profiles off Iquique. Spatial patterns of sea surface temperature and salinity from May 1996 to March 1997 followed a normal seasonal progression, though conditions were anomalously cool and fresh. Starting in March 1997, positive anomalies in sea level and sea surface temperature propagated along the South American coast to 37 degrees S. Maximum sea level anomalies occurred in two peaks in May-July 1997 and October 1997 to February 1998, separated by a relaxation period. Maximum anomalies (2 degrees C and 0.1 practical salinity units (psu)) extended to 400 m in December 1997 within 50 km of the coast. March 1998 presented the largest surface anomalies (> 4 degrees C and 0.6 psu). Strong poleward flow (20-35 cm s(-1) ) occurred to 400 m or deeper during both sea level maxima and weaker (10 cm s(-1) ) equatorward flow followed each peak. By May 1998, SST had returned to the climatological mean, and flow was equatorward next to the coast. However, offshore salinity remained anomalously high owing to a tongue of subtropical water extending southeast along the Peruvian coast. Conditions off northern Chile returned to normal between August and December 1998. The timing of the anomalies suggests a connection to equatorial waves. The progression of the 1997-1998 El Niño was very similar to that of 1982-1983, though with different timing with respect to seasons.

Bulk geochemical analysis and planktonic foraminifera of sediment core GeoB7112-5

A multivariable approach utilising bulk sediment, planktonic Foraminifera and siliceous phytoplankton has been used to reconstruct rapid variations in palaeoproductivity in the Peru-Chile Current System off northern Chile for the past 19000 cal. yr. During the early deglaciation (19000-16000 cal. yr BP), our data point to strongest upwelling intensity and highest productivity of the past 19 000 cal. yr. The late deglaciation (16000-13000 cal. yr BP) is characterised by a major change in the oceanographic setting, warmer water masses and weaker upwelling at the study site. Lowest productivity and weakest upwelling intensity are observed from the early to the middle Holocene (13000-4000 cal. yr BP), and the beginning of the late Holocene (<4000 cal. yr BP) is marked by increasing productivity, mainly driven by silicate-producing organisms. Changes in the productivity and upwelling intensity in our record may have resulted from a large-scale compression and/or displacement of the South Pacific subtropical gyre during more productive periods, in line with a northward extension of the Antarctic Circumpolar Current and increased advection of Antarctic water masses with the Peru-Chile Current. The corresponding increase in hemispheric thermal gradient and wind stress induced stronger upwelling. During the periods of lower productivity, this scenario probably reversed.

Particle flux measurements from moorings and surface sediment studies of sediment cores in the SE Pacific Ocean

Flux of siliceous plankton and taxonomic composition of diatom and silicoflagellate assemblages were determined from sediment trap samples collected in coastal upwelling-influenced waters off northern Chile (30°S, CH site) under "normal" or non-El Niño (1993-94) and El Niño conditions (1997-98). In addition, concentration of biogenic opal and siliceous plankton, and diatom and silicoflagellate assemblages preserved in surface sediments are provided for a wide area between 27° and 43°S off Chile. Regardless of the year, winter upwelling determines the maximum production pattern of siliceous microorganisms, with diatoms numerically dominating the biogenic opal flux. During the El Niño year the export is markedly lower: on an annual basis, total mass flux diminished by 60%, and diatom and silicoflagellate export by 75%. Major components of the diatom flora maintain much of their regular seasonal cycle of flux maxima and minima during both sampling periods. Neritic resting spores (RS) of Chaetoceros dominate the diatom flux, mirroring the influence of coastal-upwelled waters at the CH trap site. Occurrence of pelagic diatoms species Fragilariopsis doliolus, members of the Rhizosoleniaceae, Azpeitia spp. and Nitzschia interruptestriata, secondary components of the assemblage, reflects the intermingling of warmer waters of the Subtropical Gyre. Dictyocha messanensis dominates the silicoflagellate association almost year-around, but Distephanus pulchra delivers ca. 60% of its annual production in less than three weeks during the winter peak. The siliceous thanatocoenosis is largely dominated by diatoms, whose assemblage shows significant qualitative and quantitative variations from north to south. Between 27° and 35°S, the dominance of RS Chaetoceros, Thalassionema nitzschioides var. nitzschioides and Skeletonema costatum reflects strong export production associated with occurrence of coastal upwelling. Both highest biogenic opal content and diatom concentration at 35° and 41°-43°S coincide with highest pigment concentrations along the Chilean coast. Predominance of the diatom species Thalassiosira pacifica and T. poro-irregulata, and higher relative contribution of the silicoflagellate Distephanus speculum at 41°-43°S suggest the influence of more nutrient-rich waters and low sea surface temperatures, probably associated with the Antarctic Circumpolar Water.