Atrial septal defects can occur at various levels of the interatrial septum. Sinus venosus atrial... more Atrial septal defects can occur at various levels of the interatrial septum. Sinus venosus atrial septal defect (SVASD) results from abnormal resorption of the embryologic sinus venosus, and may be of the superior or inferior type. In this paper, we describe a 46-year-old man with inferior-type SVASD who presented with arrhythmias and dyspnea. Cardiac magnetic resonance and cardiac catheterization were useful in evaluating the anatomic/functional characteristics of inferior-type SVASD and distinguishing it from unroofed coronary sinus.
An educational program known as the Childhood Asthma Project (CAP) was implemented to reduce morb... more An educational program known as the Childhood Asthma Project (CAP) was implemented to reduce morbidity among Hispanic children with chronic asthma. Seventy-three children, ages 6-16, participated in 4 program phases: baseline assessment. one-on-one child-centered education, application, and maintenance. During baseline assessment, child and parent asthma beliefs and behaviors were evaluated and used to create educational modules on symptom recognition. peak flow meters, medications, and precipitating factors in Spanish and English. Children learned the importance of self-management, practiced using inhalers and peak flow meters and charted peak flow recordings. Videotapes provided peer modeling by showing Hispanic children with asthma performing selfmanagement tasks. During the application phase, patients practiced self-management behaviors at home and reviewed progress with a nurse educator. During maintenance, the success of self-monitoring was reviewed at follow-up appointments. Recommendations for designing health education interventions for Hispanic children are provided.
A 2-year time series (∼17-d interval) of zooplankton, sampled from June 1996 through January 1998... more A 2-year time series (∼17-d interval) of zooplankton, sampled from June 1996 through January 1998, along with oceanographic data, allowed the study of species abundances and diversity during non-El Niño and El Niño conditions at Mejillones (23° S), northern Chile. A total of 28 species was identified, of which the most abundant were the calanoids Paracalanus parvus, Calanus chilensis, Acartia tonsa, and Centropages brachiatus, the cyclopoid Oithona similis, and the poecilostomatoids Oncaea conifera and Corycaeus typicus. El Niño Southern Oscillation (El Niño) was first detected in the area in May 1997 and persisted until March 1998. We divided the time series into two parts: non-El Niño and El Niño conditions. The number of species significantly increased during El Niño conditions, but total abundance of copepods was greater during non-El Niño conditions. Although some expatriate species appeared during El Niño, such as the harpacticoid Microsetella norvegica and Eucalanus subtenuis, the diversity index of Hulburt was not significantly different between the two periods. Despite the dramatic alteration of oceanographic conditions during the El Niño period, we concluded that the community of pelagic copepods was not subject to drastic changes either in total abundance or in structure.
Deep-sea Research Part Ii-topical Studies in Oceanography, 2009
Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemi... more Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m−2 d−1 between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m−2 d−1 to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.
This study assessed the structure of the copepod community in the upper 200 m of the coastal upwe... more This study assessed the structure of the copepod community in the upper 200 m of the coastal upwelling region off central-southern Chile in late summer 2009. Vertically stratified zooplankton samples and hydrographic variables were obtained from 42 stations over the continental shelf and oceanic areas. The survey took place during active upwelling, reflected by a cold upwelling plume extending out to 150 km offshore. A total of 62 copepod species were found. Of these, Oithona similis and Paracalanusindicus accounted for ca. 60% of the whole community. Species richness (R) and the Shannon–Wiener diversity index (H′) were estimated, and the latter was additionally modified to incorporate the effect of copepod size on diversity (H′s). Samples were analyzed for two depth strata (0–50, 50–200 m) and for day vs. night conditions. Significant effects of day vs. night and strata on R, H′ and H′s indicated that diel vertical migration between these two layers was an important source of variation in the zooplankton community. H′s seemed to represent copepod diversity better than R and H′ over the spatial scale. H′s was also closely linked to colder upwelled water and the depth of the oxygen minimum zone following a principal component analysis. A positive relationship was even detected between depth of the oxygen minimum zone and H′s when strata and day/night effects were excluded. Our findings suggested that the coastal upwelling process could be an important driver of copepod diversity in this region. Upwelling leads to changes in the depth of the oxygen minimum zone and these changes impact the community composition due to species-dependent tolerances to low oxygen water.► Knowledge on controlling factors of spatial patterns of copepods in the upwelling zone is provided. ► Copepod size seems a key factor to assess spatial patterns of diversity in the upwelling zone. ► Upwelling may structure spatial diversity of copepods in the mesoscale domain. ► The oxygen minimum zone dynamics influences copepod diversity in the coastal zone.
A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from ... more A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from August 2002 to December 2003 allowed the study of annual life cycles of the copepods Calanus chilensis and Centropages brachiatus in association with environmental variability in the coastal upwelling zone. Changes in the abundance of eggs, nauplii, and copepodids were assessed from samples taken at a mean time interval of ca. 20 days. Upwelling variability in near-surface waters was reflected in seasonal changes in salinity, water column stratification, and oxycline depth, as well as a weak seasonal signal in sea surface temperature (1–2 °C). Both copepods exhibited similar life cycles, characterized by continuous reproduction throughout the year. Estimates of generation times, as a function of temperature, were 25–30 days for C. chilensis and 27–35 days for C. brachiatus, predicting about 12 and 10 generations a year, respectively. These estimates were consistent with reproduction pulses observed in the field. It was thus suggested that copepods may grow under non-limiting food conditions in this upwelling area. However, despite continuous reproduction, there were abrupt changes in population sizes along with the disappearance of early naupliar and copepodid stages taking place even during the upwelling season (spring/summer). These changes were attributed to sudden increases in mortality taking place in spring or early summer, after which the populations remained at low levels through the fall and winter. It is thus suggested that, in addition to variability in the physical environment, biological interactions modulating changes in copepod mortality should be considered for understanding copepod life cycles in highly productive upwelling systems.
Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, i... more Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, including the oceanic and coastal waters off Chile and Peru. Its annual life cycle was studied through a time-series sampling (weekly intervals) during 2002, at a fixed coastal station at an upwelling site (Mejillones Bay, 23°S) off northern Chile. The more-or-less continuous occurrence and abundance of naupliar and copepodid stages indicated that the species reproduces during most of the year, with two peaks: one during the austral spring and the other during the summer. Thereafter, an abrupt decline in the population was observed during winter. The abundances of E. inermis copepodids and nauplii were positively correlated with sea surface temperature, suggesting temperature-dependent development and growth during the spring–summer period. Three cohorts could be distinguished during one annual cycle, with generation times >30 days. The estimate of mean weight-specific daily growth (0.12 day−1) is lower than that of other species in the area, but it is consistent with a slower development rate. The distribution of this copepod is associated with the equatorial subsurface waters characterized by low-oxygen content (<0.5 ml l−1). The ascent of this water mass to the near-surface during coastal upwelling in the spring–summer period and the presence of E. inermis adults favor their reproduction and the development of cohorts in the food-rich upwelling environment. The wintertime disappearance of E. inermis from shallow waters is, thus, interpreted as a movement to deep waters (>200 m depth), probably in a lethargic mode, within the oxygen minimum zone in the adjacent oceanic area.
Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency ... more Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency of 30 days (August 2002 to December 2005), allowed the assessment of seasonal and inter-annual variation in zooplankton biomass, its C and N content, and the community structure in relation to upwelling variability. Copepods contributed 79% of the total zooplankton community and were mostly represented by Paracalanus parvus, Oithona similis, Oithona nana, Calanus chilensis, and Rhincalanus nasutus. Other copepod species, euphausiids (mainly Euphausia mucronata), gelatinous zooplankton, and crustacean larvae comprised the rest of the community. Changes in the depth of the upper boundary of the oxygen minimum zone indicated the strongly seasonal upwelling pattern. The bulk of zooplankton biomass and total copepod abundance were both strongly and positively associated with a shallow (<20 m) oxygen minimum zone; these values increased in spring/summer, when upwelling prevailed. Gelatinous zooplankton showed positive abundance anomalies in the spring and winter, whereas euphausiids had no seasonal pattern and a positive anomaly in the fall. The C content and the C/N ratio of zooplankton biomass significantly increased during the spring when chlorophyll-a was high (>5 mg m−3). No major changes in zooplankton biomass and species were found from one year to the next. We concluded that upwelling is the key process modulating variability in zooplankton biomass and its community structure in this zone. The spring/summer increase in zooplankton may be largely the result of the aggregation of dominant copepods within the upwelling region; these may reproduce throughout the year, increasing their C content and C/N ratios given high diatom concentrations.
From a time-series of sea surface temperature (SST) and zooplankton data from December 1991 throu... more From a time-series of sea surface temperature (SST) and zooplankton data from December 1991 through January 1998, and using a temperature-dependent model, we studied the growth rate (g), female size, generation time (GT), and number of generations per year of Calanus chilensis from the Mejillones Peninsula, northern Chile. Female size was negatively related to SST on both seasonal and interannual scales, and abundance was positively correlated to SST through seasons, though not associated with warm (El Niño) or cold (La Niña) years. The model predicted that cold and warm years may have a substantial effect on annual abundance of this species. Mean annual abundances of C. chilensis were not consistent with those predicted by the model, however, suggesting that factors other than temperature may regulate interannual differences in population size. For instance, changes in mortality rates between warm and cold years and advective losses under strong coastal upwelling may be implicated in the variation. Despite smaller population size in years subject to El Niño conditions, predicted annual means of g and GT showed little variability, as expected for fairly stable/constant annual mean SST. The overall g was 0.33 d 1 , while GT was 18.1 d under an overall mean SST of 17.7 C. The lack of evidence for food-shortage effects during El Niño conditions and the influence of temperature on a seasonal scale suggest that temperature-dependent growth may be a mechanism regulating population size by adjusting growth and development to stable temperature regimes on an annual basis. 2000 International Council for the Exploration of the Sea
A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out ... more A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out at the coastal upwelling zone of Mejillones (23°S, northern Chile), to analyze their annual life cycles in association with upwelling variation. These species co-exist in the upwelling zone. Weekly sampling of zooplankton and oceanographic variables including Chlorophyll-a and phytoplankton composition were obtained during January–December 2002 at a fixed station (ca. 90 m depth). Stages of abundances, their proportions, changes in body length of adult females, sex ratio and egg production rate (EPR), were used as proxies to examine copepods’ demography. Upwelling, assessed by weekly Ekman transport and oceanographic conditions, was intermittent throughout the year with lack of periodicity components. Populations of both copepod species did not correlate with these non-predictable upwelling events. C. chilensis reproduced year-round and the population showed ca. 15 peaks of adults with an average time interval between peaks of 20 days. C. brachiatus showed a similar life cycle, also having 15 peaks of adults at about 22 days of time intervals. Cross-correlation functions and spectral analysis showed that both populations correlated positively through time, but not in phase, evidencing a time lag for their reproductive cycles. The lag was also evident in their population abundances. Both species differ in their development rates and this may result in non-in phase life cycles. Our findings suggest that species-dependent attributes, such as development rates, modulated by adaptations to temperature, might impose constraints in the species life cycles determining the population cycles. Such attributes must be considered when modeling and understanding population dynamics and secondary production of copepods.
Deep-sea Research Part Ii-topical Studies in Oceanography, 2010
The copepod community structure from the Northern and Central/southern upwelling regions off Chil... more The copepod community structure from the Northern and Central/southern upwelling regions off Chile was studied and compared. The derived community descriptors were species abundance (N), species richness (R) and the Shannon-Wiener diversity index (H′). These descriptors were related to distinct habitats and conditions, sea surface temperature (SST) and depth of the upper boundary of the oxygen minimum zone (OMZ). From 159 samples, obtained between 2002 and 2008, a total number of 118 species were found of which the calanoids Paracalanus indicus, Acartia tonsa and Eucalanus inermis, along with the cyclopoid Oithona similis, and the poecilostomatoids Triconia conifera and Oncaea media were the dominant species. H′ was higher in the northern region, but no differences in N and R were detected between regions. N was higher in the epipelagic vs the deep habitat, but R and H′ did not differ. N, R and H′ correlated positively to SST and negatively to OMZ depth. The ascent of the OMZ to the upper layer forced by upwelling was proposed as a mechanism that aggregates and increases copepod diversity in the food-rich photic zone. All these findings suggest a fundamental role of upwelling variation for modulating copepod dynamics and community structure in this highly productive but strongly variable marine ecosystem.
... for Oceanographic Research in the Eastern South Pacific (COPAS), Estacion de BiologíaMarina-D... more ... for Oceanographic Research in the Eastern South Pacific (COPAS), Estacion de BiologíaMarina-Dichato, Departamento ... were obtained from a time series study developed by the Universidad de Antofagasta and ... with 4 terminal setae, of which 3 are long and 1 lateral seta short. ...
In the Humboldt Current System, the region off central-southern Chile has relatively high eddy ki... more In the Humboldt Current System, the region off central-southern Chile has relatively high eddy kinetic energy, generating an extensive coastal transition zone (∼600 km offshore) in which coastally derived eddies are recurrent features. This energy might promote strong exchanges of water, biogeochemical properties, and plankton between the coastal upwelling band and the adjacent oceanic zone. In this study, the mesoscale structure of epipelagic copepod assemblages and its relationship to environmental variability and the eddy field in the coastal transition zone and oceanic areas off Concepción (34–39°S, 73–84°W) were investigated. Zooplankton samples were collected during cruises of opportunity carried out during the 2006 coastal upwelling season and the oceanographic conditions were derived from satellite data on sea surface height, temperature, and chlorophyll a. The use of cluster analyses and indicator species revealed two main copepod assemblages: (i) species with a mainly coastal distribution, the maximum in total abundance being found nearshore and (ii) species with a mainly oceanic distribution (beyond ∼500 km from the coast), where a secondary maximum was observed. Both types of assemblages, however, included species widely distributed in the coastal transition zone. An ordination analysis identified sea surface temperature and chlorophyll a as the main factors affecting the distribution of most species and the clustering of stations resembled the distribution of these variables. Some of the large calanoid species that are common in shelf waters were more abundant within nearshore eddies rich in chlorophyll a and other cyclonic eddies far offshore; however, the eddy field alone did not explain the copepod mesoscale distribution. Altogether, the wide distribution of shelf/slope copepod species in this region suggests that physical and biological mechanisms might be acting to extend the productive area of the coastal upwelling zone.
Journal of Experimental Marine Biology and Ecology
Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL... more Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL) and histology of gonads have been investigated with Calanus chilensis females sampled weekly, from late November to December 2004, at a station located in the coastal zone off Dichato (Chile), at time diatom concentration in phytoplankton bloom was high. Weekly EPR estimate in nature did not change significantly during this period. It remained close to normal values (25-40 eggs/female/day), whereas HS was constantly low and high proportions of AL were observed. In parallel, bioassays revealed that EPR was strongly depressed by artificially enriched diets, corresponding to natural diatom assemblages (NDA) occurring in the field, while abnormal HS and AL values could not be improved. Ingestion of diatoms by females was estimated by faecal pellet production rates and SEM examination of diatom remains in pellet samples. Low HS and the high amounts of abnormal larvae were not reversible when females were offered a favourable food, the dinoflagellate P. minimum (PM). Minor cell degradations were observed in gonads of females fed NDA diets. In comparison with other environments, present results show that impairment of Calanoid copepod reproductive factors can occur at both high and low diatom concentrations, depending on maternal diets and diatom species in blooms.
A field experiment was carried out in October 1998 during active upwelling in a coastal area off ... more A field experiment was carried out in October 1998 during active upwelling in a coastal area off the Mejillones Peninsula (23°S). Zooplankton was sampled at day and night, during two subsequent days at 4 stations inside and outside of the upwelling plume. Three depth strata were sampled: 0–20 m, 20–80 m and 80–200 m. Oceanographic data were obtained in a grid of 23 stations using a CTDO, a fluorometer and a Doppler current meter. Calanus chilensis was mostly represented by late stages, i.e. copepodid C5 and adult males and females. There were no day/night effects on vertical distribution, and abundance was significantly higher inside the upwelling plume in the upper 20-m layer at nearly 14 ind. m−3, compared to ca. 5 ind. m−3 outside the upwelling plume. Temperature at 10 m depth and biomass, estimated from stage numbers and their mean dry weights, were used to estimate growth and daily production of Calanus at temperature-dependent rates. The potential loss of biomass from the upwelling center because of advection in the upwelling plume was estimated from current data in the Ekman layer and biomass density. The mean cross-shelf component of the current was estimated at 10.4km d−1 within the upwelling plume. This yielded a loss of biomass of 9.7 mg dry weight m−2. Production, estimated by a temperature-dependent approach, ranged between 44 and 35 mg dry weight m−2 d−1, at mean temperatures of 14.6°C and 15.8°C inside and outside of the upwelling plume respectively. Within the plume, as much as 22% of daily production may be advected offshore. However, a higher concentration of biomass in the upwelling plume allowed a greater production compared to surrounding areas. A mass balance approach suggests that advective losses may not have a major impact on the C. chilensis population, because of very high daily production at temperature-dependent rates.
Journal of Experimental Marine Biology and Ecology, 2007
Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL... more Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL) and histology of gonads have been investigated with Calanus chilensis females sampled weekly, from late November to December 2004, at a station located in the coastal zone off Dichato (Chile), at time diatom concentration in phytoplankton bloom was high. Weekly EPR estimate in nature did not change significantly during this period. It remained close to normal values (25–40 eggs/female/day), whereas HS was constantly low and high proportions of AL were observed. In parallel, bioassays revealed that EPR was strongly depressed by artificially enriched diets, corresponding to natural diatom assemblages (NDA) occurring in the field, while abnormal HS and AL values could not be improved. Ingestion of diatoms by females was estimated by faecal pellet production rates and SEM examination of diatom remains in pellet samples. Low HS and the high amounts of abnormal larvae were not reversible when females were offered a favourable food, the dinoflagellate P. minimum (PM). Minor cell degradations were observed in gonads of females fed NDA diets. In comparison with other environments, present results show that impairment of Calanoid copepod reproductive factors can occur at both high and low diatom concentrations, depending on maternal diets and diatom species in blooms.
A 2-year time series (∼ 17-d interval) of zooplankton, sampled from June 1996 through January 199... more A 2-year time series (∼ 17-d interval) of zooplankton, sampled from June 1996 through January 1998, along with oceanographic data, allowed the study of species abundances and diversity during non-El Niño and El Niño conditions at Mejillones (23° S) northern Chile. A total of 28 species was identified, of which the most abundant were the calanoids Paracalanus parvus, Calanus chilensis, Acartia tonsa, and Centropages brachiatus, the cyclopoid Oithona similis, and the poecilostomatoids Oncaea conifera and Corycaeus typicus. El Niño Southern Oscillation (El Niño) was first detected in the area in May 1997 and persisted until March 1998. We divided the time series into two parts: non-El Niño conditions. The number of species significantly increased during Eland El Niño conditions, but total abundance of copepods was greater during non-El Niño conditions. Although some expatriate species appeared during El Niño, such as the harpacticoid Microsetella norvegica and Eucalanus subtenuis, the diversity index of Hulburt was not significantly different between the two periods. Despite the dramatic alteration of oceanographic conditions during the El Niño period, we concluded that the community of pelagic copepods was not subject to drastic changes either in total abundance or in structure.
Atrial septal defects can occur at various levels of the interatrial septum. Sinus venosus atrial... more Atrial septal defects can occur at various levels of the interatrial septum. Sinus venosus atrial septal defect (SVASD) results from abnormal resorption of the embryologic sinus venosus, and may be of the superior or inferior type. In this paper, we describe a 46-year-old man with inferior-type SVASD who presented with arrhythmias and dyspnea. Cardiac magnetic resonance and cardiac catheterization were useful in evaluating the anatomic/functional characteristics of inferior-type SVASD and distinguishing it from unroofed coronary sinus.
An educational program known as the Childhood Asthma Project (CAP) was implemented to reduce morb... more An educational program known as the Childhood Asthma Project (CAP) was implemented to reduce morbidity among Hispanic children with chronic asthma. Seventy-three children, ages 6-16, participated in 4 program phases: baseline assessment. one-on-one child-centered education, application, and maintenance. During baseline assessment, child and parent asthma beliefs and behaviors were evaluated and used to create educational modules on symptom recognition. peak flow meters, medications, and precipitating factors in Spanish and English. Children learned the importance of self-management, practiced using inhalers and peak flow meters and charted peak flow recordings. Videotapes provided peer modeling by showing Hispanic children with asthma performing selfmanagement tasks. During the application phase, patients practiced self-management behaviors at home and reviewed progress with a nurse educator. During maintenance, the success of self-monitoring was reviewed at follow-up appointments. Recommendations for designing health education interventions for Hispanic children are provided.
A 2-year time series (∼17-d interval) of zooplankton, sampled from June 1996 through January 1998... more A 2-year time series (∼17-d interval) of zooplankton, sampled from June 1996 through January 1998, along with oceanographic data, allowed the study of species abundances and diversity during non-El Niño and El Niño conditions at Mejillones (23° S), northern Chile. A total of 28 species was identified, of which the most abundant were the calanoids Paracalanus parvus, Calanus chilensis, Acartia tonsa, and Centropages brachiatus, the cyclopoid Oithona similis, and the poecilostomatoids Oncaea conifera and Corycaeus typicus. El Niño Southern Oscillation (El Niño) was first detected in the area in May 1997 and persisted until March 1998. We divided the time series into two parts: non-El Niño and El Niño conditions. The number of species significantly increased during El Niño conditions, but total abundance of copepods was greater during non-El Niño conditions. Although some expatriate species appeared during El Niño, such as the harpacticoid Microsetella norvegica and Eucalanus subtenuis, the diversity index of Hulburt was not significantly different between the two periods. Despite the dramatic alteration of oceanographic conditions during the El Niño period, we concluded that the community of pelagic copepods was not subject to drastic changes either in total abundance or in structure.
Deep-sea Research Part Ii-topical Studies in Oceanography, 2009
Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemi... more Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m−2 d−1 between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m−2 d−1 to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.
This study assessed the structure of the copepod community in the upper 200 m of the coastal upwe... more This study assessed the structure of the copepod community in the upper 200 m of the coastal upwelling region off central-southern Chile in late summer 2009. Vertically stratified zooplankton samples and hydrographic variables were obtained from 42 stations over the continental shelf and oceanic areas. The survey took place during active upwelling, reflected by a cold upwelling plume extending out to 150 km offshore. A total of 62 copepod species were found. Of these, Oithona similis and Paracalanusindicus accounted for ca. 60% of the whole community. Species richness (R) and the Shannon–Wiener diversity index (H′) were estimated, and the latter was additionally modified to incorporate the effect of copepod size on diversity (H′s). Samples were analyzed for two depth strata (0–50, 50–200 m) and for day vs. night conditions. Significant effects of day vs. night and strata on R, H′ and H′s indicated that diel vertical migration between these two layers was an important source of variation in the zooplankton community. H′s seemed to represent copepod diversity better than R and H′ over the spatial scale. H′s was also closely linked to colder upwelled water and the depth of the oxygen minimum zone following a principal component analysis. A positive relationship was even detected between depth of the oxygen minimum zone and H′s when strata and day/night effects were excluded. Our findings suggested that the coastal upwelling process could be an important driver of copepod diversity in this region. Upwelling leads to changes in the depth of the oxygen minimum zone and these changes impact the community composition due to species-dependent tolerances to low oxygen water.► Knowledge on controlling factors of spatial patterns of copepods in the upwelling zone is provided. ► Copepod size seems a key factor to assess spatial patterns of diversity in the upwelling zone. ► Upwelling may structure spatial diversity of copepods in the mesoscale domain. ► The oxygen minimum zone dynamics influences copepod diversity in the coastal zone.
A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from ... more A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from August 2002 to December 2003 allowed the study of annual life cycles of the copepods Calanus chilensis and Centropages brachiatus in association with environmental variability in the coastal upwelling zone. Changes in the abundance of eggs, nauplii, and copepodids were assessed from samples taken at a mean time interval of ca. 20 days. Upwelling variability in near-surface waters was reflected in seasonal changes in salinity, water column stratification, and oxycline depth, as well as a weak seasonal signal in sea surface temperature (1–2 °C). Both copepods exhibited similar life cycles, characterized by continuous reproduction throughout the year. Estimates of generation times, as a function of temperature, were 25–30 days for C. chilensis and 27–35 days for C. brachiatus, predicting about 12 and 10 generations a year, respectively. These estimates were consistent with reproduction pulses observed in the field. It was thus suggested that copepods may grow under non-limiting food conditions in this upwelling area. However, despite continuous reproduction, there were abrupt changes in population sizes along with the disappearance of early naupliar and copepodid stages taking place even during the upwelling season (spring/summer). These changes were attributed to sudden increases in mortality taking place in spring or early summer, after which the populations remained at low levels through the fall and winter. It is thus suggested that, in addition to variability in the physical environment, biological interactions modulating changes in copepod mortality should be considered for understanding copepod life cycles in highly productive upwelling systems.
Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, i... more Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, including the oceanic and coastal waters off Chile and Peru. Its annual life cycle was studied through a time-series sampling (weekly intervals) during 2002, at a fixed coastal station at an upwelling site (Mejillones Bay, 23°S) off northern Chile. The more-or-less continuous occurrence and abundance of naupliar and copepodid stages indicated that the species reproduces during most of the year, with two peaks: one during the austral spring and the other during the summer. Thereafter, an abrupt decline in the population was observed during winter. The abundances of E. inermis copepodids and nauplii were positively correlated with sea surface temperature, suggesting temperature-dependent development and growth during the spring–summer period. Three cohorts could be distinguished during one annual cycle, with generation times >30 days. The estimate of mean weight-specific daily growth (0.12 day−1) is lower than that of other species in the area, but it is consistent with a slower development rate. The distribution of this copepod is associated with the equatorial subsurface waters characterized by low-oxygen content (<0.5 ml l−1). The ascent of this water mass to the near-surface during coastal upwelling in the spring–summer period and the presence of E. inermis adults favor their reproduction and the development of cohorts in the food-rich upwelling environment. The wintertime disappearance of E. inermis from shallow waters is, thus, interpreted as a movement to deep waters (>200 m depth), probably in a lethargic mode, within the oxygen minimum zone in the adjacent oceanic area.
Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency ... more Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency of 30 days (August 2002 to December 2005), allowed the assessment of seasonal and inter-annual variation in zooplankton biomass, its C and N content, and the community structure in relation to upwelling variability. Copepods contributed 79% of the total zooplankton community and were mostly represented by Paracalanus parvus, Oithona similis, Oithona nana, Calanus chilensis, and Rhincalanus nasutus. Other copepod species, euphausiids (mainly Euphausia mucronata), gelatinous zooplankton, and crustacean larvae comprised the rest of the community. Changes in the depth of the upper boundary of the oxygen minimum zone indicated the strongly seasonal upwelling pattern. The bulk of zooplankton biomass and total copepod abundance were both strongly and positively associated with a shallow (<20 m) oxygen minimum zone; these values increased in spring/summer, when upwelling prevailed. Gelatinous zooplankton showed positive abundance anomalies in the spring and winter, whereas euphausiids had no seasonal pattern and a positive anomaly in the fall. The C content and the C/N ratio of zooplankton biomass significantly increased during the spring when chlorophyll-a was high (>5 mg m−3). No major changes in zooplankton biomass and species were found from one year to the next. We concluded that upwelling is the key process modulating variability in zooplankton biomass and its community structure in this zone. The spring/summer increase in zooplankton may be largely the result of the aggregation of dominant copepods within the upwelling region; these may reproduce throughout the year, increasing their C content and C/N ratios given high diatom concentrations.
From a time-series of sea surface temperature (SST) and zooplankton data from December 1991 throu... more From a time-series of sea surface temperature (SST) and zooplankton data from December 1991 through January 1998, and using a temperature-dependent model, we studied the growth rate (g), female size, generation time (GT), and number of generations per year of Calanus chilensis from the Mejillones Peninsula, northern Chile. Female size was negatively related to SST on both seasonal and interannual scales, and abundance was positively correlated to SST through seasons, though not associated with warm (El Niño) or cold (La Niña) years. The model predicted that cold and warm years may have a substantial effect on annual abundance of this species. Mean annual abundances of C. chilensis were not consistent with those predicted by the model, however, suggesting that factors other than temperature may regulate interannual differences in population size. For instance, changes in mortality rates between warm and cold years and advective losses under strong coastal upwelling may be implicated in the variation. Despite smaller population size in years subject to El Niño conditions, predicted annual means of g and GT showed little variability, as expected for fairly stable/constant annual mean SST. The overall g was 0.33 d 1 , while GT was 18.1 d under an overall mean SST of 17.7 C. The lack of evidence for food-shortage effects during El Niño conditions and the influence of temperature on a seasonal scale suggest that temperature-dependent growth may be a mechanism regulating population size by adjusting growth and development to stable temperature regimes on an annual basis. 2000 International Council for the Exploration of the Sea
A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out ... more A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out at the coastal upwelling zone of Mejillones (23°S, northern Chile), to analyze their annual life cycles in association with upwelling variation. These species co-exist in the upwelling zone. Weekly sampling of zooplankton and oceanographic variables including Chlorophyll-a and phytoplankton composition were obtained during January–December 2002 at a fixed station (ca. 90 m depth). Stages of abundances, their proportions, changes in body length of adult females, sex ratio and egg production rate (EPR), were used as proxies to examine copepods’ demography. Upwelling, assessed by weekly Ekman transport and oceanographic conditions, was intermittent throughout the year with lack of periodicity components. Populations of both copepod species did not correlate with these non-predictable upwelling events. C. chilensis reproduced year-round and the population showed ca. 15 peaks of adults with an average time interval between peaks of 20 days. C. brachiatus showed a similar life cycle, also having 15 peaks of adults at about 22 days of time intervals. Cross-correlation functions and spectral analysis showed that both populations correlated positively through time, but not in phase, evidencing a time lag for their reproductive cycles. The lag was also evident in their population abundances. Both species differ in their development rates and this may result in non-in phase life cycles. Our findings suggest that species-dependent attributes, such as development rates, modulated by adaptations to temperature, might impose constraints in the species life cycles determining the population cycles. Such attributes must be considered when modeling and understanding population dynamics and secondary production of copepods.
Deep-sea Research Part Ii-topical Studies in Oceanography, 2010
The copepod community structure from the Northern and Central/southern upwelling regions off Chil... more The copepod community structure from the Northern and Central/southern upwelling regions off Chile was studied and compared. The derived community descriptors were species abundance (N), species richness (R) and the Shannon-Wiener diversity index (H′). These descriptors were related to distinct habitats and conditions, sea surface temperature (SST) and depth of the upper boundary of the oxygen minimum zone (OMZ). From 159 samples, obtained between 2002 and 2008, a total number of 118 species were found of which the calanoids Paracalanus indicus, Acartia tonsa and Eucalanus inermis, along with the cyclopoid Oithona similis, and the poecilostomatoids Triconia conifera and Oncaea media were the dominant species. H′ was higher in the northern region, but no differences in N and R were detected between regions. N was higher in the epipelagic vs the deep habitat, but R and H′ did not differ. N, R and H′ correlated positively to SST and negatively to OMZ depth. The ascent of the OMZ to the upper layer forced by upwelling was proposed as a mechanism that aggregates and increases copepod diversity in the food-rich photic zone. All these findings suggest a fundamental role of upwelling variation for modulating copepod dynamics and community structure in this highly productive but strongly variable marine ecosystem.
... for Oceanographic Research in the Eastern South Pacific (COPAS), Estacion de BiologíaMarina-D... more ... for Oceanographic Research in the Eastern South Pacific (COPAS), Estacion de BiologíaMarina-Dichato, Departamento ... were obtained from a time series study developed by the Universidad de Antofagasta and ... with 4 terminal setae, of which 3 are long and 1 lateral seta short. ...
In the Humboldt Current System, the region off central-southern Chile has relatively high eddy ki... more In the Humboldt Current System, the region off central-southern Chile has relatively high eddy kinetic energy, generating an extensive coastal transition zone (∼600 km offshore) in which coastally derived eddies are recurrent features. This energy might promote strong exchanges of water, biogeochemical properties, and plankton between the coastal upwelling band and the adjacent oceanic zone. In this study, the mesoscale structure of epipelagic copepod assemblages and its relationship to environmental variability and the eddy field in the coastal transition zone and oceanic areas off Concepción (34–39°S, 73–84°W) were investigated. Zooplankton samples were collected during cruises of opportunity carried out during the 2006 coastal upwelling season and the oceanographic conditions were derived from satellite data on sea surface height, temperature, and chlorophyll a. The use of cluster analyses and indicator species revealed two main copepod assemblages: (i) species with a mainly coastal distribution, the maximum in total abundance being found nearshore and (ii) species with a mainly oceanic distribution (beyond ∼500 km from the coast), where a secondary maximum was observed. Both types of assemblages, however, included species widely distributed in the coastal transition zone. An ordination analysis identified sea surface temperature and chlorophyll a as the main factors affecting the distribution of most species and the clustering of stations resembled the distribution of these variables. Some of the large calanoid species that are common in shelf waters were more abundant within nearshore eddies rich in chlorophyll a and other cyclonic eddies far offshore; however, the eddy field alone did not explain the copepod mesoscale distribution. Altogether, the wide distribution of shelf/slope copepod species in this region suggests that physical and biological mechanisms might be acting to extend the productive area of the coastal upwelling zone.
Journal of Experimental Marine Biology and Ecology
Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL... more Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL) and histology of gonads have been investigated with Calanus chilensis females sampled weekly, from late November to December 2004, at a station located in the coastal zone off Dichato (Chile), at time diatom concentration in phytoplankton bloom was high. Weekly EPR estimate in nature did not change significantly during this period. It remained close to normal values (25-40 eggs/female/day), whereas HS was constantly low and high proportions of AL were observed. In parallel, bioassays revealed that EPR was strongly depressed by artificially enriched diets, corresponding to natural diatom assemblages (NDA) occurring in the field, while abnormal HS and AL values could not be improved. Ingestion of diatoms by females was estimated by faecal pellet production rates and SEM examination of diatom remains in pellet samples. Low HS and the high amounts of abnormal larvae were not reversible when females were offered a favourable food, the dinoflagellate P. minimum (PM). Minor cell degradations were observed in gonads of females fed NDA diets. In comparison with other environments, present results show that impairment of Calanoid copepod reproductive factors can occur at both high and low diatom concentrations, depending on maternal diets and diatom species in blooms.
A field experiment was carried out in October 1998 during active upwelling in a coastal area off ... more A field experiment was carried out in October 1998 during active upwelling in a coastal area off the Mejillones Peninsula (23°S). Zooplankton was sampled at day and night, during two subsequent days at 4 stations inside and outside of the upwelling plume. Three depth strata were sampled: 0–20 m, 20–80 m and 80–200 m. Oceanographic data were obtained in a grid of 23 stations using a CTDO, a fluorometer and a Doppler current meter. Calanus chilensis was mostly represented by late stages, i.e. copepodid C5 and adult males and females. There were no day/night effects on vertical distribution, and abundance was significantly higher inside the upwelling plume in the upper 20-m layer at nearly 14 ind. m−3, compared to ca. 5 ind. m−3 outside the upwelling plume. Temperature at 10 m depth and biomass, estimated from stage numbers and their mean dry weights, were used to estimate growth and daily production of Calanus at temperature-dependent rates. The potential loss of biomass from the upwelling center because of advection in the upwelling plume was estimated from current data in the Ekman layer and biomass density. The mean cross-shelf component of the current was estimated at 10.4km d−1 within the upwelling plume. This yielded a loss of biomass of 9.7 mg dry weight m−2. Production, estimated by a temperature-dependent approach, ranged between 44 and 35 mg dry weight m−2 d−1, at mean temperatures of 14.6°C and 15.8°C inside and outside of the upwelling plume respectively. Within the plume, as much as 22% of daily production may be advected offshore. However, a higher concentration of biomass in the upwelling plume allowed a greater production compared to surrounding areas. A mass balance approach suggests that advective losses may not have a major impact on the C. chilensis population, because of very high daily production at temperature-dependent rates.
Journal of Experimental Marine Biology and Ecology, 2007
Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL... more Variations of egg production rate (EPR), hatching success (HS), production of abnormal larvae (AL) and histology of gonads have been investigated with Calanus chilensis females sampled weekly, from late November to December 2004, at a station located in the coastal zone off Dichato (Chile), at time diatom concentration in phytoplankton bloom was high. Weekly EPR estimate in nature did not change significantly during this period. It remained close to normal values (25–40 eggs/female/day), whereas HS was constantly low and high proportions of AL were observed. In parallel, bioassays revealed that EPR was strongly depressed by artificially enriched diets, corresponding to natural diatom assemblages (NDA) occurring in the field, while abnormal HS and AL values could not be improved. Ingestion of diatoms by females was estimated by faecal pellet production rates and SEM examination of diatom remains in pellet samples. Low HS and the high amounts of abnormal larvae were not reversible when females were offered a favourable food, the dinoflagellate P. minimum (PM). Minor cell degradations were observed in gonads of females fed NDA diets. In comparison with other environments, present results show that impairment of Calanoid copepod reproductive factors can occur at both high and low diatom concentrations, depending on maternal diets and diatom species in blooms.
A 2-year time series (∼ 17-d interval) of zooplankton, sampled from June 1996 through January 199... more A 2-year time series (∼ 17-d interval) of zooplankton, sampled from June 1996 through January 1998, along with oceanographic data, allowed the study of species abundances and diversity during non-El Niño and El Niño conditions at Mejillones (23° S) northern Chile. A total of 28 species was identified, of which the most abundant were the calanoids Paracalanus parvus, Calanus chilensis, Acartia tonsa, and Centropages brachiatus, the cyclopoid Oithona similis, and the poecilostomatoids Oncaea conifera and Corycaeus typicus. El Niño Southern Oscillation (El Niño) was first detected in the area in May 1997 and persisted until March 1998. We divided the time series into two parts: non-El Niño conditions. The number of species significantly increased during Eland El Niño conditions, but total abundance of copepods was greater during non-El Niño conditions. Although some expatriate species appeared during El Niño, such as the harpacticoid Microsetella norvegica and Eucalanus subtenuis, the diversity index of Hulburt was not significantly different between the two periods. Despite the dramatic alteration of oceanographic conditions during the El Niño period, we concluded that the community of pelagic copepods was not subject to drastic changes either in total abundance or in structure.
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Papers by Pamela Hidalgo