Kendra L. Daly

An index to assess the health and benefits of the global ocean

The ocean plays a critical role in supporting human well-being, from providing food, livelihoods and recreational opportunities to regulating the global climate. Sustainable management aimed at maintaining the flow of a broad range of benefits from the ocean requires a comprehensive and quantitative method to measure and monitor the health of coupled human–ocean systems. We created an index comprising ten diverse public goals for a healthy coupled human–ocean system and calculated the index for every coastal country. Globally, the overall index score was 60 out of 100 (range 36–86), with developed countries generally performing better than developing countries, but with notable exceptions. Only 5% of countries scored higher than 70, whereas 32% scored lower than 50. The index provides a powerful tool to raise public awareness, direct resource management, improve policy and prioritize scientific research.

Abundance and distribution of krill in the ice edge zone of the Weddell Sea, austral spring 1983

Abstract The effect of the ice edge zone on the abundance and distribution of Antarctic krill was investigated by acoustic and net sampling methods in the northwestern Weddell Sea during austral spring, 1983. The dominant acoustic target was Euphausia superba which occurred over the entire study area in the upper 100 m of the water column, either in aggregations or loosely defined layers. The vertical depth distribution under the pack ice was similar to that of open water. Average krill biomass under the ice ranged from 1 to 68 g m −2 and in open water range from 10 to 100 g m −2 . Another euphausiid, Thysanoessa macrura , appeared to be ubiquitous but was much less abundant than E. superba. T. macrura also formed aggregations. All stages were found in open water, but only adults and juveniles under the ice. A few adults and larvae of a third euphausiid, E. frigida , were found only in open water. Salps, which were abundant in open water, were not present under the pack ice. The pack ice directly influenced the abundance and distribution of young krill. Juvenile E. superba were much more abundant under the ice than in open water north of the ice edge. Numerous observations were made of juvenile krill feeding on the undersurface of ice floes where chlorophyll concentrations were much greater than in the water column below. These observations appear to indicate that the pack ice serves as a giant nursery area for young krill, providing a refugium from predators as well as a vast food resource that may be critical to the survival of overwintering juvenile E. superba .

Non‐Redfield carbon and nitrogen cycling in the Arctic: Effects of ecosystem structure and dynamics

The C:N ratio is a critical parameter used in both global ocean carbon models and field studies to understand carbon and nutrient cycling as well as to estimate exported carbon from the euphotic zone. The so-called Redfield ratio (C:N = 6.6 by atoms) [Redfield et al., 1963] is widely used for such calculations. Here we present data from the NE Greenland continental shelf that show that most of the C:N ratios for particulate (autotrophic and heterotrophic) and dissolved pools and rates of transformation among them exceed Redfield proportions from June to August, owing to species composition, size, and biological interactions. The ecosystem components that likely comprised sinking particles and had relatively high C:N ratios (geometric means) included (1) the particulate organic matter (C:N = 8.9) dominated by nutrient-deficient diatoms, resulting from low initial nitrate concentrations (approximately 4 μM) in Arctic surface waters; (2) the dominant zooplankton, herbivorous copepods (C:N = 9.6), having lipid storage typical of Arctic copepods; and (3) copepod fecal pellets (C:N = 33.2). Relatively high dissolved organic carbon concentrations (median 105 μM) were approximately 25 to 45 μM higher than reported for other systems and may be broadly characteristic of Arctic waters. A carbon-rich dissolved organic carbon pool also was generated during summer. Since the magnitude of carbon and nitrogen uncoupling in the surface mixed layer appeared to be greater than in other regions and occurred throughout the productive season, the C:N ratio of particulate organic matter may be a better conversion factor than the Redfield ratio to estimate carbon export for broad application in northern high-latitude systems.

Toxicity and mutagenicity of Gulf of Mexico waters during and after the deepwater horizon oil spill.

The Deepwater Horizon oil spill is unparalleled among environmental hydrocarbon releases, because of the tremendous volume of oil, the additional contamination by dispersant, and the oceanic depth at which this release occurred. Here, we present data on general toxicity and mutagenicity of upper water column waters and, to a lesser degree, sediment porewater of the Northeastern Gulf of Mexico (NEGOM) and west Florida shelf (WFS) at the time of the Deepwater Horizon oil spill in 2010 and thereafter. During a research cruise in August 2010, analysis of water collected in the NEGOM indicated that samples of 3 of 14 (21%) stations were toxic to bacteria based on the Microtox assay, 4 of 13 (34%) were toxic to phytoplankton via the QwikLite assay, and 6 of 14 (43%) showed DNA damaging activity using the λ-Microscreen Prophage induction assay. The Microtox and Microscreen assays indicated that the degree of toxicity was correlated to total petroleum hydrocarbon concentration. Long-term monitoring of stations on the NEGOM and the WFS was undertaken by 8 and 6 cruises to these areas, respectively. Microtox toxicity was nearly totally absent by December 2010 in the Northeastern Gulf of Mexico (3 of 8 cruises with one positive station). In contrast, QwikLite toxicity assay yielded positives at each cruise, often at multiple stations or depths, indicating the greater sensitivity of the QwikLite assay to environmental factors. The Microscreen mutagenicity assays indicated that certain water column samples overlying the WFS were mutagenic at least 1.5 years after capping the Macondo well. Similarly, sediment porewater samples taken from 1000, 1200, and 1400 m from the slope off the WFS in June 2011 were also highly genotoxic. Our observations are consistent with a portion of the dispersed oil from the Macondo well area advecting to the southeast and upwelling onto the WFS, although other explanations exist. Organisms in contact with these waters might experience DNA damage that could lead to mutation and heritable alterations to the community pangenome. Such mutagenic interactions might not become apparent in higher organisms for years.

New production in the Northeast Water Polynya: 1993

The Northeast Water Polynya has been suggested as acting as a sink for carbon, especially during the spring and summer when phytoplankton growth is active. During 1993 the polynya was sampled for the entire growing period (late May through mid-August) in order to more accurately assess the magnitude, controls and patterns of new and total (ammonium, nitrate and urea) nitrogen production. This represents the first assessment of new production throughout an entire season in the Arctic. We found that, in 1993, new production, based on 15N-tracer techniques and integrated over the euphotic zone, was 0.141 mmol N m−2 h−1 (0.361 g C m−2 d−1 when converted using observed C/N ratios). Measured f ratios averaged 0.65 and demonstrate that the system, to a great extent, was using nitrate as a nitrogen source. In general f ratios were greatest early in the season and minimal in mid-summer. Urea uptake was highly variable and contributed slightly less than ammonium to phytoplankton nitrogen demand. Nitrate uptake at stations with low (< 0.5μM) nitrate concentrations was significantly reduced, implying that nitrate concentrations limited phytoplankton growth late in the growing season. Long-term new production rates calculated from nutrient depletion patterns from the polynya as a whole as well as a time-series constructed from a single location were ca. 0.144-0.281 g C m−2 d−1. The relationship between new production as measured by incubations and nutrient depletion budgets suggests that phytoplankton growth is the dominant factor influencing the nitrogen budget of the polynya. The amount of material available for removal from the euphotic zone is limited and constrains the degree to which the polynya can act as a regional carbon sink.

Flux of particulate matter through copepods in the Northeast water polynya

Particulate organic carbon (POC) and nitrogen (PON) production by large calanoid copepods was investigated on the northeast Greenland shelf during August 1992 and May to August 1993. Both Calanus hyperboreus and C. glacialis females, when suspended in seawater collected from the chlorophyll maximum, produced about 40 pellets per day, which contained a carbon and nitrogen content equivalent to 8% and 6% of body carbon, respectively, and 2% of body nitrogen. In experiments, the carbon:nitrogen (C:N) ratio by weight of suspended particulates, C. hyperboreus, and fecal pellets was 6.7, 7.7 and 28.5, respectively. The unusually high C:N ratio for pellets, in part, may be attributed to elevated ratios of > 20μm size fractions of particulate organic matter, the size fraction more common in the diet of these large copepods and the fraction dominated by diatoms according to microscopic and pigment data. The implied elevated C:N ratios of large phytoplankton cells were probably due to nitrogen deficiency, as shown by other studies in this region. In addition, female C. hyperboreus appeared to be more efficient in assimilating nitrogen than carbon, which also would have contributed to high C:N ratios in egested pellets. Unfractionated POC concentrations explained 54% of the variability in carbon egestion and 70% of the variability in nitrogen egestion in copepods, whereas copepod body content accounted for little of the variation on the short time scales of the experiments. Carbon egestion by C. hyperboreus was positively correlated with POC concentrations at the depth of the chlorophyll maximum, while nitrogen egestion was negatively correlated with PON concentrations in the euphotic zone. Estimates of potential community egestion rates for the upper water column indicate that copepods represent a major pathway of organic carbon transformation in this Arctic shelf system. On average, copepods may have ingested 45% of the primary production and egested fecal matter equivalent to 20% of the carbon and 12% of the nitrogen particulate flux sedimenting from the surface layer. However, several lines of evidence suggest that pellets were remineralized in the water column and, hence, may have contributed little organic carbon and nitrogen to the benthos.

Particulate Dimethylsulfoniopropionate Removal and Dimethylsulfide Production by Zooplankton in the Southern Ocean

The influence of Antarctic krill, Euphausia superba, on particulate dimethlysulfoniopropionate (DMSP(p)) and dimethylsulfide (DMS) concentrations in surface waters of the Southern Ocean was investigated by shipboard experiments during austral spring near the Antarctic Peninsula. Chlorophyll concentrations were low in the water column, but substantially higher in sea ice due to the high biomass of ice algae, predominantly pennate diatoms. A comparison of DMSP(p) concentrations and algal accessory pigments indicated that DMSP(p) was associated primarily with diatoms (fucoxanthin) and to a minor extent with Phaeocystis spp. (19’-hexanoyloxyfucoxanthin) in sea ice algae. Maximum DMSP(p) and fucoxanthin concentrations also occurred in the 100-200 μm size fraction. We interpret this to mean that high biomass of diatoms in sea ice contributes significantly to DMSP(p) pools in the Antarctic.

Acoustic scattering from zooplankton and micronekton in relation to a whale feeding site near Georges Bank and Cape Cod

Abstract This research was part of the South Channel Ocean Productivity Experiment (SCOPEX), a multidisciplinary study to investigate the biological and physical processes associated with the very high annual springtime abundance of right whales ( Eubalaena glacialis ) in the Great South Channel off New England. Right whales appear to gather there in the spring because of the increased abundance of aggregations of their principal prey, the copepod Calanus finmarchicus . Observations of hydroacoustic scattering were made in relation to the hydrography, whale distributions, and other biological measurements in the vicinity of the Great South Channel during May 1986, March, April and May of 1988, and May and June of 1989. Copepods were detected (at 200 kHz) as a near-surface layer with strong diel changes. In 1989, a second frequency (120 kHz) was used to discriminate between copepod layers (which the 120 kHz detected only weakly) and other targets (which both frequencies detected). Acoustically distinct layers of zooplankton and micronekton were observed, which were often correlated in time and space with the copepod layers. Quantitative estimates derived from the acoustic data indicate that the abundance of zooplankton varied from 1–5 g wet weight m −3 to 18–25 g wet weight m −3 which correlates well with the abundances observed from MOCNESS tows. The acoustic data revealed a complex diel migration of two layers in addition to the copepods. Euphausiids (predominantly Meganyctiphanes sp.) were found in a layer above the bottom, and a mid-water layer may have been due to sand lance ( Ammodytes americanus ). The observed biological phenomena appeared to be related to the complex hydrography of the region. A surface thermal front existed at the northern entrance to the channel in 1988 and 1989, with colder vertically mixed water to the south and warmer stratified water to the north. A Fast Fourier Transform analysis for spectral composition and autocovariance shows (a) strong contrasts in the spectral density across one frontal feature (predominantly a salinity front) as opposed to away from the front, and (b) significant differences between those areas where a whale moved more rapidly (presumably searching for food) and where it spent more time (presumably or observably feeding). The behavior of whales, in particular the right whale, can be shown to be related to the spatial scales and abundance of their prey by the use of hydroacoustic estimates of target distribution and abundance.

Hydrography, nutrients, and carbon pools in the Pacific sector of the Southern Ocean: Implications for carbon flux

We investigated the hydrography, nutrients, and dissolved and particulate carbon pools in the western Pacific sector of the Antarctic Circumpolar Current (ACC) during austral summer 1996 to assess the regions role in the carbon cycle. Low fCO2 values along two transects indicated that much of the study area was a sink for atmospheric CO2. The fCO2 values were lowest near the Polar Front (PF) and the Subtropical Front (STF), concomitant with maxima of chlorophyll a and particulate and dissolved organic carbon. The largest biomass accumulations did not occur at fronts, which had high surface geostrophic velocities (20–51 cm s−1), but in relatively low velocity regions near fronts or in an eddy. Thus vertical motion and horizontal advection associated with fronts may have replenished nutrients in surface waters but also dispersed phytoplankton. Although surface waters north of the PF have been characterized as a “high nutrient-low chlorophyll” region, low silicic acid (Si) concentrations (2–4 μM) may limit production of large diatoms and therefore the potential carbon flux. Low concentrations (4–10 μM Si) at depths of winter mixing constrain the level of Si replenishment to surface waters. It has been suggested that an increase in aeolian iron north of the PF may increase primary productivity and carbon export. Our results, however, indicate that while diatom growth and carbon export may be enhanced, the extent ultimately would be limited by the vertical supply of Si. South of the PF, the primary mechanism by which carbon is exported to deep water appears to be through diatom flux. We suggest that north of the PF, particulate and dissolved carbon may be exported primarily to intermediate depths through subduction and diapycnal mixing associated with Subantarctic Mode Water and Antarctic Intermediate Water formation. These physical-biological interactions and Si dynamics should be included in future biogeochemical models to provide a more accurate prediction of carbon flux.

Mechanisms of Patch Formation

Many mechanisms both physical (e.g., light, temperature, ocean currents, density gradients, topography) and biological (e.g., allelopathy, competition, predation, selective foraging) are considered responsible for patch formation. Wiens (1976) presented an excellent review of population responses to environmental patchiness. He identified localized random disturbances (e.g., fire, erosion, tree windfalls), predation, selective herbivory, and vegetational patterns as potential causes of patch formation. Roughgarden (1977) discussed five general mechanisms that are responsible for patchiness: resource distribution, dispersal, aggregation behavior, competition, and reaction-diffusion.