... Patricia Ayón Dejo, Katia Aronés Flores. ... During El Niño 1972-1973, sardine, light fish, j... more ... Patricia Ayón Dejo, Katia Aronés Flores. ... During El Niño 1972-1973, sardine, light fish, jack mackerel, flat fish and cutlassfish (Sandoval de Castillo, 1985) were the main species, while during El Niño 1982-1983 species including anchovy, sardine, mackerel (eg jack and bullet ...
The Northern Humboldt Current system (NHCS) produces more fish per unit area than any other regio... more The Northern Humboldt Current system (NHCS) produces more fish per unit area than any other region in the world. Although the system produces enough macrozooplankton to sustain its high production of forage fish, the paucity of information on macrozooplankton hampers research into the system. In this study, we estimated the biomass of the epipelagic crustacean macrozooplankton from the NHCS during both austral summer and spring 2005. To do this, we developed a bi-frequency acoustic method and extracted high-resolution information on the biomass and the patterns of distribution of crustacean macrozooplankton, fish and other marine compartments. We found that, although macrozooplankton comprises a number of distinct organisms, the euphausiids were the zooplankton group that better fitted the patterns from independent net sampling zooplankton data. Also, the similarities between the nocturnal patterns of size and biomass macrozooplankton distribution from this study and the known patterns of euphausiids, in particular Euphausia mucronata, suggest that euphausiids were the main constituent of the estimated nocturnal acoustic macrozooplankton biomass even if other organisms such as large copepods may have contributed considerably to the macrozooplankton biomass. The total macrozooplankton biomass was estimated to about 105 g m−2, i.e., two to five times more than previous estimates. This direct biomass estimation of macrozooplankton is in agreement with the new findings in trophic ecology indicating that forage fish consume mainly macrozooplankton. This high biomass also supports the current hypotheses explaining the NHCS high fish production. Using the method, we are able to revisit present-day and historical acoustic databases and extract high-resolution data on macrozooplankton, a key ecological compartment of the ecosystem. Since zooplankton is the link between the physically driven primary producers and the biologically driven tertiary consumers, this information is essential to achieve a mechanistic understanding of the system, from physics to top predators.► Information on crustacean macrozooplankton and fish was extracted from acoustic data. ► Euphausiids were the macrozooplankton group that best fitted the acoustic patterns. ► The total macrozooplankton biomass was estimated to about 105 g m−2. ► This biomass estimation is in agreement with new findings in trophic ecology. ► It also supports current hypotheses explaining the system’s high fish production.
Journal of The European Optical Society-rapid Publications, 2010
Zooplankton is a key element in aquatic food webs. Rapid mapping of abundance, combined with info... more Zooplankton is a key element in aquatic food webs. Rapid mapping of abundance, combined with information on taxonomic and size composition is necessary to understand ecosystem dynamics. Classical sampling with towed plankton nets does not allow resolving fine scale distributions along hydrographic gradients (e.g. fronts and clines) although such structures determine community assemblages and trophic interactions. Furthermore, sample analysis is labor intensive and time consuming. To overcome these shortcomings, Lightframe On-sight Keyspecies Investigation (LOKI), a new imaging device, was developed for sensing spatial variability of plankton distribution on scales below the 1 m level. Here, we give a brief description of the LOKI system and demonstrate its potential for taxonomic identification using images of various zooplankton taxa collected in the south east Pacific.
The objective of the present study was to verify possible spatial, seasonal, and inter-annual cha... more The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.
Planktonic ostracods are deep dwelling mesozooplankton that are diel migrators and detritus feede... more Planktonic ostracods are deep dwelling mesozooplankton that are diel migrators and detritus feeders. In the Humboldt Current region off Peru they are shown to be strongly influenced by El Niño conditions. Species composition and distribution were compared in day and night (vertically hauled 50–0 m Hensen Net) samples that were collected from three latitudinal zones (3°40' S–6°S; 9°S–12°S and 15°S–18°S) during 1998 (El Niño year) and in 2001 (a year with usual upwelling regime). Unexpectedly, ostracods diversity and abundance were much higher during El Niño than during the upwelling regime. Tropical species that normally occurred north of 6°S, spread southwards and increased in abundance; Euconchoecia aculeata was often dominant, particularly in coastal waters, and the frequency of occurrence of others increased throughout the entire region (e.g. Porroecia porrecta, Archiconchoecia striata). Conversely, Conchoecetta giesbrechti, a species associated with subsurface oxygen minimum waters, was more frequently found and in higher abundance during the upwelling regime in 2001. Thus, El Niño conditions apparently favor the occupation of the epipelagic zone by many halocyprid species advected in from tropical waters, possibly because the higher oxygen content in the subsurface waters enables these diel migrating species to survive better. Additional factors may be a consequence of the switch to a picoplankton production cycle, which results in more detritus being retained in the upper waters, and the sharp reduction in planktivorous fishes such as the anchovy which may reduce the predation pressure they experience.
A review of zooplankton studies conducted in Peruvian marine waters is given. After a short histo... more A review of zooplankton studies conducted in Peruvian marine waters is given. After a short history of the development of zooplankton research off Peru, we review zooplankton methodology, taxonomy, biodiversity, spatial distribution, seasonal and interannual variability, trophodynamics, secondary production, and modelling. We review studies on several micro-, meso-, macro-, and meroplankton groups, and give a species list from both published and unpublished reports. Three regional zooplankton groups have been identified: (1) a continental shelf group dominated by Acartia tonsa and Centropages brachiatus; (2) a continental slope group characterized by siphonophores, bivalves, foraminifera and radiolaria; (3) and a species-rich oceanic group. The highest zooplankton abundances and biomasses were often found between 4–6°S and 14–16°S, where continental shelves are narrow. Species composition changes with distance from the shore. Species composition and biomass also vary strongly on short time scales due to advection, peaks of larval production, trophic interactions, and community succession. The relation of zooplankton to climatic variability (ENSO and multi-decadal) and fish stocks is discussed in the context of ecological regime shifts. An intermediate upwelling hypothesis is proposed, based on the negative effects of low upwelling intensity in summer or extremely strong and enduring winter upwelling on zooplankton abundance off Peru. According to this hypothesis, intermediate upwelling creates an optimal environmental window for zooplankton communities. Finally, we highlight important knowledge gaps that warrant attention in future.
Vertical distribution and diel vertical migration of a zooplankton community were studied at two ... more Vertical distribution and diel vertical migration of a zooplankton community were studied at two stations off Central Peru in April 2006. Zooplankton was collected at five depth strata by vertical hauls with Hydo-Bios multinet (300-μm mesh, 0.25-m2 mouth size). The zooplankton community was distributed in relation to a strong, shallow oxycline (1 ml l−1 oxygen isopleth generally above 36 m). The highest total abundance was always in the upper, well-oxygenated layer. The most important species were: Acartia tonsa (72.86%), Centropages brachiatus (7.5%), and Paracalanus parvus (3.1%); Acartia tonsa was the dominant species at all times. Larvae of the polychaete Magelona sp. (7.5%) and larvae of the brachiopod Discinisca lamellosa (3.5%) were numerically dominant in April and small copepods e.g. Oncaea venusta (3.88%) were numerically dominant during August. Five distinct patterns of vertical distribution and migration in relation to the oxygen minimum layer were distinguished in this study: (1) Ontogenetic vertical migration through the oxycline (Acartia tonsa adults, nauplii, and copepodids), (2) permanent limitation to layers above the oxycline (e.g. Oikopleura sp., most invertebrate larvae), (3) distribution mostly below the oxycline with occasional migration into the layers just above the oxycline (Eucalanus inermis), (4) Diel Vertical Migration (Centropages brachiatus), and (5) reverse Diel Vertical Migration (larvae of the polychaete Magelona sp.).
The El Niño of 1997-98 was one of the strongest warming events of the past century; among many ot... more The El Niño of 1997-98 was one of the strongest warming events of the past century; among many other effects, it impacted phytoplankton along the Peruvian coast by changing species composition and reducing biomass. While responses of the main fish resources to this natural perturbation are relatively well known, understanding the ecosystem response as a whole requires an ecotrophic multispecies approach. In this work, we construct trophic models of the Northern Humboldt Current Ecosystem (NHCE) and compare the La Niña (LN) years in 1995-96 with the El Niño (EN) years in 1997-98. The model area extends from 4°S-16°S and to 60 nm from the coast. The model consists of 32 functional groups of organisms and differs from previous trophic models of the Peruvian system through: (i) division of plankton into size classes to account for EN-associated changes and feeding preferences of small pelagic fish, (ii) increased division of demersal groups and separation of life history stages of hake, (iii) inclusion of mesopelagic fish, and (iv) incorporation of the jumbo squid (Dosidicus gigas), which became abundant following EN. Results show that EN reduced the size and organization of energy flows of the NHCE, but the overall functioning (proportion of energy flows used for respiration, consumption by predators, detritus and export) of the ecosystem was maintained. The reduction of diatom biomass during EN forced omnivorous planktivorous fish to switch to a more zooplankton-dominated diet, raising their trophic level. Consequently, in the EN model the trophic level increased for several predatory groups (mackerel, other large pelagics, sea birds, pinnipeds) and for fishery catch. A high modeled biomass of macrozooplankton was needed to balance the consumption by planktivores, especially during EN condition when observed diatoms biomass diminished dramatically. Despite overall lower planktivorous fish catches, the higher primary production required-to-catch ratio implied a stronger ecological impact of the fishery and stresses the need for precautionary management of fisheries during and after EN. During EN energetic indicators such as the lower primary production/total biomass ratio suggest a more energetically efficient ecosystem, while reduced network indicators such as the cycling index and relative ascendency indicate of a less organized state of the ecosystem. Compared to previous trophic models of the NHCE we observed: (i) a shrinking of ecosystem size in term of energy flows, (ii) slight changes in overall functioning (proportion of energy flows used for respiration, consumption by predators and detritus), and (iii) the use of alternate pathways leading to a higher ecological impact of the fishery for planktivorous fish.
The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feed... more The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feeding fish, which suggested that their population dynamics may ‘drive’ or ‘control’ ecosystem dynamics. With this in mind we analysed the relationship between forage fish populations and their main prey, zooplankton populations. Our study combined a zooplankton sampling program (1961–2005) with simultaneous acoustic observations on fish from 40 pelagic surveys (1983–2005) conducted by the Peruvian Marine Research Institute (IMARPE) and landing statistics for anchoveta (Engraulis ringens) and sardine (Sardinops sagax) along the Peruvian coast from 1961 to 2005. The multi-year trend of anchoveta population abundance varied consistently with zooplankton biovolume trend, suggesting bottom-up control on anchovy at the population scale (since oceanographic conditions and phytoplankton production support the changes in zooplankton abundance). For a finer-scale analysis (km) we statistically modelled zooplankton biovolume as a function of geographical (latitude and distance from the 200-m isobath), environmental (sea surface temperature), temporal (year, month and time-of-day) and biological (acoustic anchovy and sardine biomass within 5 km of each zooplankton sample) covariates over all survey using both classification and regression trees (CART) and generalized additive models (GAM). CART showed local anchoveta density to have the strongest effect on zooplankton biovolume, with significantly reduced levels of biovolume for higher neighbourhood anchoveta biomass. Additionally, zooplankton biovolume was higher offshore than on the shelf. GAM results corroborated the CART findings, also showing a clear diel effect on zooplankton biovolume, probably due to diel migration or daytime net avoidance. Apparently, the observed multi-year population scale bottom-up control is not inconsistent with local depletion of zooplankton when anchoveta are locally abundant, since the latter effect was observed over a wide range of overall anchoveta abundance.
We used an individual-based model of anchovy (Engraulis ringens) early life history coupled with ... more We used an individual-based model of anchovy (Engraulis ringens) early life history coupled with hydrodynamic outputs from the regional oceanic modeling system (ROMS) to investigate the factors driving variability in egg and larval survival rates in the northern Humboldt upwelling region off Peru. Individuals were released within a coastal area and followed for a period of 30 days. Those that were still in the coastal area at that time were considered as retained. We investigated the spatial and temporal variability in the release locations of the individuals retained, and compared these to observed egg concentration patterns reconstructed from a 40-year period of monitoring. A first set of simulations using passive particles to represent anchovy eggs and larvae revealed a large sensitivity of the results to the initial vertical distribution of particles. We then conducted two additional sets of simulations that included the effect of egg buoyancy, larval vertical swimming behavior and lethal temperature. We obtained (1) maximal coastal retention close to the surface in winter and in deeper layers in summer, (2) a large influence of egg buoyancy and of larval vertical behavior on coastal retention in all seasons, (3) a partial match between dates and locations of enhanced retention and observed egg concentration patterns and (4) a low effect of lethal temperature on survival except when associated with high egg density. The model suggests that an optimal temporal spawning pattern for maximizing coastal retention would have two maximums, the most significant in austral winter and the second in summer. This pattern agrees roughly with observed spawning seasonality, but with temporal discrepancy of about two months in the peaks of both series. Spatially, we obtained higher retention from 10 S to 20 S, whereas the observed maximum egg concentration was located between 6°S and 14°S. Among the three sets of simulations, the one taking into account larval vertical swimming behavior lead to the best match with the data.
The Northern Humboldt Current Ecosystem is one of the most productive in the world in terms of fi... more The Northern Humboldt Current Ecosystem is one of the most productive in the world in terms of fish production. Its location near to the equator permits strong upwelling under relatively low winds, thus creating optimal conditions for the development of plankton communities. These communities ultimately support abundant populations of grazing fish such as the Peruvian anchoveta, Engraulis ringens. The ecosystem is also subject to strong inter-annual environmental variability associated with the El Niño Southern Oscillation (ENSO), which has major effects on nutrient structure, primary production, and higher trophic levels. Here our objective is to model the contributions of several external drivers (i.e. reconstructed phytoplankton changes, fish immigration, and fishing rate) and internal control mechanisms (i.e. predator-prey) to ecosystem dynamics over an ENSO cycle. Steady-state models and timeseries data from the Instituto del Mar del Perú (IMARPE) from 1995 to 2004 provide the base data for simulations conducted with the program Ecopath with Ecosim. In simulations all three external drivers contribute to ecosystem dynamics. Changes in phytoplankton quantity and composition (i.e. contribution of diatoms and dino-and silicoflagellates), as affected by upwelling intensity, were important in dynamics of the El Niño of 1997-98 and the subsequent 3 years. The expansion and immigration of mesopelagic fish populations during El Niño was important for dynamics in following years. Fishing rate changes were the most important of the three external drivers tested, helping to explain observed dynamics throughout the modeled period, and particularly during the post-El Niño period. Internal control settings show a mix of predator-prey control settings; however a ''wasp-waist" control of the ecosystem by small pelagic fish is not supported.
A Lagrangian model is used to simulate and quantify in the northern Humboldt upwelling ecosystem ... more A Lagrangian model is used to simulate and quantify in the northern Humboldt upwelling ecosystem the processes of enrichment, concentration and retention, identified by Bakun [Bakun, A., 1996. Patterns in the ocean. Ocean processes and marine population dynamics. University of California Sea Grant, California, USA, in cooperation with Centro de Investigaciones Biologicas de Noroeste, La Paz, Baja California Sur, Mexico, 323 pp.] as being important for the survival and recruitment of early life stages of pelagic fish. The method relies on tracking the positions of particles within water velocity fields generated by a three-dimensional hydrodynamic model. Simple criteria for considering particles as participating to enrichment, concentration or retention are used to derive indices of the three processes. We analyse the spatial distribution of and seasonal variability in these indices. The results are discussed in relation to anchovy (Engraulis ringens) eggs and larvae distributions off Peru, and to a comparable study conducted in the southern Benguela upwelling ecosystem.
... Patricia Ayón Dejo, Katia Aronés Flores. ... During El Niño 1972-1973, sardine, light fish, j... more ... Patricia Ayón Dejo, Katia Aronés Flores. ... During El Niño 1972-1973, sardine, light fish, jack mackerel, flat fish and cutlassfish (Sandoval de Castillo, 1985) were the main species, while during El Niño 1982-1983 species including anchovy, sardine, mackerel (eg jack and bullet ...
The Northern Humboldt Current system (NHCS) produces more fish per unit area than any other regio... more The Northern Humboldt Current system (NHCS) produces more fish per unit area than any other region in the world. Although the system produces enough macrozooplankton to sustain its high production of forage fish, the paucity of information on macrozooplankton hampers research into the system. In this study, we estimated the biomass of the epipelagic crustacean macrozooplankton from the NHCS during both austral summer and spring 2005. To do this, we developed a bi-frequency acoustic method and extracted high-resolution information on the biomass and the patterns of distribution of crustacean macrozooplankton, fish and other marine compartments. We found that, although macrozooplankton comprises a number of distinct organisms, the euphausiids were the zooplankton group that better fitted the patterns from independent net sampling zooplankton data. Also, the similarities between the nocturnal patterns of size and biomass macrozooplankton distribution from this study and the known patterns of euphausiids, in particular Euphausia mucronata, suggest that euphausiids were the main constituent of the estimated nocturnal acoustic macrozooplankton biomass even if other organisms such as large copepods may have contributed considerably to the macrozooplankton biomass. The total macrozooplankton biomass was estimated to about 105 g m−2, i.e., two to five times more than previous estimates. This direct biomass estimation of macrozooplankton is in agreement with the new findings in trophic ecology indicating that forage fish consume mainly macrozooplankton. This high biomass also supports the current hypotheses explaining the NHCS high fish production. Using the method, we are able to revisit present-day and historical acoustic databases and extract high-resolution data on macrozooplankton, a key ecological compartment of the ecosystem. Since zooplankton is the link between the physically driven primary producers and the biologically driven tertiary consumers, this information is essential to achieve a mechanistic understanding of the system, from physics to top predators.► Information on crustacean macrozooplankton and fish was extracted from acoustic data. ► Euphausiids were the macrozooplankton group that best fitted the acoustic patterns. ► The total macrozooplankton biomass was estimated to about 105 g m−2. ► This biomass estimation is in agreement with new findings in trophic ecology. ► It also supports current hypotheses explaining the system’s high fish production.
Journal of The European Optical Society-rapid Publications, 2010
Zooplankton is a key element in aquatic food webs. Rapid mapping of abundance, combined with info... more Zooplankton is a key element in aquatic food webs. Rapid mapping of abundance, combined with information on taxonomic and size composition is necessary to understand ecosystem dynamics. Classical sampling with towed plankton nets does not allow resolving fine scale distributions along hydrographic gradients (e.g. fronts and clines) although such structures determine community assemblages and trophic interactions. Furthermore, sample analysis is labor intensive and time consuming. To overcome these shortcomings, Lightframe On-sight Keyspecies Investigation (LOKI), a new imaging device, was developed for sensing spatial variability of plankton distribution on scales below the 1 m level. Here, we give a brief description of the LOKI system and demonstrate its potential for taxonomic identification using images of various zooplankton taxa collected in the south east Pacific.
The objective of the present study was to verify possible spatial, seasonal, and inter-annual cha... more The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.
Planktonic ostracods are deep dwelling mesozooplankton that are diel migrators and detritus feede... more Planktonic ostracods are deep dwelling mesozooplankton that are diel migrators and detritus feeders. In the Humboldt Current region off Peru they are shown to be strongly influenced by El Niño conditions. Species composition and distribution were compared in day and night (vertically hauled 50–0 m Hensen Net) samples that were collected from three latitudinal zones (3°40' S–6°S; 9°S–12°S and 15°S–18°S) during 1998 (El Niño year) and in 2001 (a year with usual upwelling regime). Unexpectedly, ostracods diversity and abundance were much higher during El Niño than during the upwelling regime. Tropical species that normally occurred north of 6°S, spread southwards and increased in abundance; Euconchoecia aculeata was often dominant, particularly in coastal waters, and the frequency of occurrence of others increased throughout the entire region (e.g. Porroecia porrecta, Archiconchoecia striata). Conversely, Conchoecetta giesbrechti, a species associated with subsurface oxygen minimum waters, was more frequently found and in higher abundance during the upwelling regime in 2001. Thus, El Niño conditions apparently favor the occupation of the epipelagic zone by many halocyprid species advected in from tropical waters, possibly because the higher oxygen content in the subsurface waters enables these diel migrating species to survive better. Additional factors may be a consequence of the switch to a picoplankton production cycle, which results in more detritus being retained in the upper waters, and the sharp reduction in planktivorous fishes such as the anchovy which may reduce the predation pressure they experience.
A review of zooplankton studies conducted in Peruvian marine waters is given. After a short histo... more A review of zooplankton studies conducted in Peruvian marine waters is given. After a short history of the development of zooplankton research off Peru, we review zooplankton methodology, taxonomy, biodiversity, spatial distribution, seasonal and interannual variability, trophodynamics, secondary production, and modelling. We review studies on several micro-, meso-, macro-, and meroplankton groups, and give a species list from both published and unpublished reports. Three regional zooplankton groups have been identified: (1) a continental shelf group dominated by Acartia tonsa and Centropages brachiatus; (2) a continental slope group characterized by siphonophores, bivalves, foraminifera and radiolaria; (3) and a species-rich oceanic group. The highest zooplankton abundances and biomasses were often found between 4–6°S and 14–16°S, where continental shelves are narrow. Species composition changes with distance from the shore. Species composition and biomass also vary strongly on short time scales due to advection, peaks of larval production, trophic interactions, and community succession. The relation of zooplankton to climatic variability (ENSO and multi-decadal) and fish stocks is discussed in the context of ecological regime shifts. An intermediate upwelling hypothesis is proposed, based on the negative effects of low upwelling intensity in summer or extremely strong and enduring winter upwelling on zooplankton abundance off Peru. According to this hypothesis, intermediate upwelling creates an optimal environmental window for zooplankton communities. Finally, we highlight important knowledge gaps that warrant attention in future.
Vertical distribution and diel vertical migration of a zooplankton community were studied at two ... more Vertical distribution and diel vertical migration of a zooplankton community were studied at two stations off Central Peru in April 2006. Zooplankton was collected at five depth strata by vertical hauls with Hydo-Bios multinet (300-μm mesh, 0.25-m2 mouth size). The zooplankton community was distributed in relation to a strong, shallow oxycline (1 ml l−1 oxygen isopleth generally above 36 m). The highest total abundance was always in the upper, well-oxygenated layer. The most important species were: Acartia tonsa (72.86%), Centropages brachiatus (7.5%), and Paracalanus parvus (3.1%); Acartia tonsa was the dominant species at all times. Larvae of the polychaete Magelona sp. (7.5%) and larvae of the brachiopod Discinisca lamellosa (3.5%) were numerically dominant in April and small copepods e.g. Oncaea venusta (3.88%) were numerically dominant during August. Five distinct patterns of vertical distribution and migration in relation to the oxygen minimum layer were distinguished in this study: (1) Ontogenetic vertical migration through the oxycline (Acartia tonsa adults, nauplii, and copepodids), (2) permanent limitation to layers above the oxycline (e.g. Oikopleura sp., most invertebrate larvae), (3) distribution mostly below the oxycline with occasional migration into the layers just above the oxycline (Eucalanus inermis), (4) Diel Vertical Migration (Centropages brachiatus), and (5) reverse Diel Vertical Migration (larvae of the polychaete Magelona sp.).
The El Niño of 1997-98 was one of the strongest warming events of the past century; among many ot... more The El Niño of 1997-98 was one of the strongest warming events of the past century; among many other effects, it impacted phytoplankton along the Peruvian coast by changing species composition and reducing biomass. While responses of the main fish resources to this natural perturbation are relatively well known, understanding the ecosystem response as a whole requires an ecotrophic multispecies approach. In this work, we construct trophic models of the Northern Humboldt Current Ecosystem (NHCE) and compare the La Niña (LN) years in 1995-96 with the El Niño (EN) years in 1997-98. The model area extends from 4°S-16°S and to 60 nm from the coast. The model consists of 32 functional groups of organisms and differs from previous trophic models of the Peruvian system through: (i) division of plankton into size classes to account for EN-associated changes and feeding preferences of small pelagic fish, (ii) increased division of demersal groups and separation of life history stages of hake, (iii) inclusion of mesopelagic fish, and (iv) incorporation of the jumbo squid (Dosidicus gigas), which became abundant following EN. Results show that EN reduced the size and organization of energy flows of the NHCE, but the overall functioning (proportion of energy flows used for respiration, consumption by predators, detritus and export) of the ecosystem was maintained. The reduction of diatom biomass during EN forced omnivorous planktivorous fish to switch to a more zooplankton-dominated diet, raising their trophic level. Consequently, in the EN model the trophic level increased for several predatory groups (mackerel, other large pelagics, sea birds, pinnipeds) and for fishery catch. A high modeled biomass of macrozooplankton was needed to balance the consumption by planktivores, especially during EN condition when observed diatoms biomass diminished dramatically. Despite overall lower planktivorous fish catches, the higher primary production required-to-catch ratio implied a stronger ecological impact of the fishery and stresses the need for precautionary management of fisheries during and after EN. During EN energetic indicators such as the lower primary production/total biomass ratio suggest a more energetically efficient ecosystem, while reduced network indicators such as the cycling index and relative ascendency indicate of a less organized state of the ecosystem. Compared to previous trophic models of the NHCE we observed: (i) a shrinking of ecosystem size in term of energy flows, (ii) slight changes in overall functioning (proportion of energy flows used for respiration, consumption by predators and detritus), and (iii) the use of alternate pathways leading to a higher ecological impact of the fishery for planktivorous fish.
The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feed... more The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feeding fish, which suggested that their population dynamics may ‘drive’ or ‘control’ ecosystem dynamics. With this in mind we analysed the relationship between forage fish populations and their main prey, zooplankton populations. Our study combined a zooplankton sampling program (1961–2005) with simultaneous acoustic observations on fish from 40 pelagic surveys (1983–2005) conducted by the Peruvian Marine Research Institute (IMARPE) and landing statistics for anchoveta (Engraulis ringens) and sardine (Sardinops sagax) along the Peruvian coast from 1961 to 2005. The multi-year trend of anchoveta population abundance varied consistently with zooplankton biovolume trend, suggesting bottom-up control on anchovy at the population scale (since oceanographic conditions and phytoplankton production support the changes in zooplankton abundance). For a finer-scale analysis (km) we statistically modelled zooplankton biovolume as a function of geographical (latitude and distance from the 200-m isobath), environmental (sea surface temperature), temporal (year, month and time-of-day) and biological (acoustic anchovy and sardine biomass within 5 km of each zooplankton sample) covariates over all survey using both classification and regression trees (CART) and generalized additive models (GAM). CART showed local anchoveta density to have the strongest effect on zooplankton biovolume, with significantly reduced levels of biovolume for higher neighbourhood anchoveta biomass. Additionally, zooplankton biovolume was higher offshore than on the shelf. GAM results corroborated the CART findings, also showing a clear diel effect on zooplankton biovolume, probably due to diel migration or daytime net avoidance. Apparently, the observed multi-year population scale bottom-up control is not inconsistent with local depletion of zooplankton when anchoveta are locally abundant, since the latter effect was observed over a wide range of overall anchoveta abundance.
We used an individual-based model of anchovy (Engraulis ringens) early life history coupled with ... more We used an individual-based model of anchovy (Engraulis ringens) early life history coupled with hydrodynamic outputs from the regional oceanic modeling system (ROMS) to investigate the factors driving variability in egg and larval survival rates in the northern Humboldt upwelling region off Peru. Individuals were released within a coastal area and followed for a period of 30 days. Those that were still in the coastal area at that time were considered as retained. We investigated the spatial and temporal variability in the release locations of the individuals retained, and compared these to observed egg concentration patterns reconstructed from a 40-year period of monitoring. A first set of simulations using passive particles to represent anchovy eggs and larvae revealed a large sensitivity of the results to the initial vertical distribution of particles. We then conducted two additional sets of simulations that included the effect of egg buoyancy, larval vertical swimming behavior and lethal temperature. We obtained (1) maximal coastal retention close to the surface in winter and in deeper layers in summer, (2) a large influence of egg buoyancy and of larval vertical behavior on coastal retention in all seasons, (3) a partial match between dates and locations of enhanced retention and observed egg concentration patterns and (4) a low effect of lethal temperature on survival except when associated with high egg density. The model suggests that an optimal temporal spawning pattern for maximizing coastal retention would have two maximums, the most significant in austral winter and the second in summer. This pattern agrees roughly with observed spawning seasonality, but with temporal discrepancy of about two months in the peaks of both series. Spatially, we obtained higher retention from 10 S to 20 S, whereas the observed maximum egg concentration was located between 6°S and 14°S. Among the three sets of simulations, the one taking into account larval vertical swimming behavior lead to the best match with the data.
The Northern Humboldt Current Ecosystem is one of the most productive in the world in terms of fi... more The Northern Humboldt Current Ecosystem is one of the most productive in the world in terms of fish production. Its location near to the equator permits strong upwelling under relatively low winds, thus creating optimal conditions for the development of plankton communities. These communities ultimately support abundant populations of grazing fish such as the Peruvian anchoveta, Engraulis ringens. The ecosystem is also subject to strong inter-annual environmental variability associated with the El Niño Southern Oscillation (ENSO), which has major effects on nutrient structure, primary production, and higher trophic levels. Here our objective is to model the contributions of several external drivers (i.e. reconstructed phytoplankton changes, fish immigration, and fishing rate) and internal control mechanisms (i.e. predator-prey) to ecosystem dynamics over an ENSO cycle. Steady-state models and timeseries data from the Instituto del Mar del Perú (IMARPE) from 1995 to 2004 provide the base data for simulations conducted with the program Ecopath with Ecosim. In simulations all three external drivers contribute to ecosystem dynamics. Changes in phytoplankton quantity and composition (i.e. contribution of diatoms and dino-and silicoflagellates), as affected by upwelling intensity, were important in dynamics of the El Niño of 1997-98 and the subsequent 3 years. The expansion and immigration of mesopelagic fish populations during El Niño was important for dynamics in following years. Fishing rate changes were the most important of the three external drivers tested, helping to explain observed dynamics throughout the modeled period, and particularly during the post-El Niño period. Internal control settings show a mix of predator-prey control settings; however a ''wasp-waist" control of the ecosystem by small pelagic fish is not supported.
A Lagrangian model is used to simulate and quantify in the northern Humboldt upwelling ecosystem ... more A Lagrangian model is used to simulate and quantify in the northern Humboldt upwelling ecosystem the processes of enrichment, concentration and retention, identified by Bakun [Bakun, A., 1996. Patterns in the ocean. Ocean processes and marine population dynamics. University of California Sea Grant, California, USA, in cooperation with Centro de Investigaciones Biologicas de Noroeste, La Paz, Baja California Sur, Mexico, 323 pp.] as being important for the survival and recruitment of early life stages of pelagic fish. The method relies on tracking the positions of particles within water velocity fields generated by a three-dimensional hydrodynamic model. Simple criteria for considering particles as participating to enrichment, concentration or retention are used to derive indices of the three processes. We analyse the spatial distribution of and seasonal variability in these indices. The results are discussed in relation to anchovy (Engraulis ringens) eggs and larvae distributions off Peru, and to a comparable study conducted in the southern Benguela upwelling ecosystem.
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Papers by Patricia Ayon