Blue carbon gains from glacial retreat along Antarctic fjords: What should we expect?

Rising atmospheric CO₂ is intensifying climate change but it is also driving global and particularly polar greening. However, most blue carbon sinks (that held by marine organisms) are shrinking, which is important as these are hotspots of genuine carbon sequestration. Polar blue carbon increases with losses of marine ice over high latitude continental shelf areas. Marine ice (sea ice, ice shelf and glacier retreat) losses generate a valuable negative feedback on climate change. Blue carbon change with sea ice and ice shelf losses has been estimated, but not how blue carbon responds to glacier retreat along fjords. We derive a testable estimate of glacier retreat driven blue carbon gains by investigating three fjords in the West Antarctic Peninsula (WAP). We started by multiplying ~40 year mean glacier retreat rates by the number of retreating WAP fjords and their time of exposure. We multiplied this area by regional zoobenthic carbon means from existing datasets to suggest that WAP fjords generate 3,130 tonnes of new zoobenthic carbon per year (t zC/year) and sequester >780 t zC/year. We tested this by capture and analysis of 204 high resolution seabed images along emerging WAP fjords. Biota within these images were identified to density per 13 functional groups. Mean stored carbon per individual was assigned from literature values to give a stored zoobenthic Carbon per area, which was multiplied up by area of fjord exposed over time, which increased the estimate to 4,536 t zC/year. The purpose of this study was to establish a testable estimate of blue carbon change caused by glacier retreat along Antarctic fjords and thus to establish its relative importance compared to polar and other carbon sinks.     

Perspective: Increasing blue carbon around Antarctica is an ecosystem service of considerable societal and economic value worth protecting

Precautionary conservation and cooperative global governance are needed to protect Antarctic blue carbon: the world's largest increasing natural form of carbon storage with high sequestration potential. As patterns of ice loss around Antarctica become more uniform, there is an underlying increase in carbon capture‐to‐storage‐to‐sequestration on the seafloor. The amount of carbon captured per unit area is increasing and the area available to blue carbon is also increasing. Carbon sequestration could further increase under moderate (+1°C) ocean warming, contrary to decreasing global blue carbon stocks elsewhere. For example, in warmer waters, mangroves and seagrasses are in decline and benthic organisms are close to their physiological limits, so a 1°C increase in water temperature could push them above their thermal tolerance (e.g. bleaching of coral reefs). In contrast, on the basis of past change and current research, we expect that Antarctic blue carbon could increase by orders of magnitude. The Antarctic seafloor is biophysically unique and the site of carbon sequestration, the benthos, faces less anthropogenic disturbance than any other ocean continental shelf environment. This isolation imparts both vulnerability to change, and an avenue to conserve one of the world's last biodiversity refuges. In economic terms, the value of Antarctic blue carbon is estimated at between £0.65 and £1.76 billion (~2.27 billion USD) for sequestered carbon in the benthos around the continental shelf. To balance biodiversity protection against society's economic objectives, this paper builds on a proposal incentivising protection by building a ‘non‐market framework’ via the 2015 Paris Agreement to the United Nations Framework Convention on Climate Change. This could be connected and coordinated through the Antarctic Treaty System to promote and motivate member states to value Antarctic blue carbon and maintain scientific integrity and conservation for the positive societal values ingrained in the Antarctic Treaty System.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

Polar zoobenthos blue carbon storage increases with sea ice losses,because across‐shelf growth gains from longer algal blooms outweigh ice scour mortality in the shallows

One of the major climate‐forced global changes has been white to blue to green; losses of sea ice extent in time and space around Arctic and West Antarctic seas has increased open water and the duration (though not magnitude) of phytoplankton blooms. Blueing of the poles has increases potential for heat absorption for positive feedback but conversely the longer phytoplankton blooms have increased carbon export to storage and sequestration by shelf benthos. However, ice shelf collapses and glacier retreat can calve more icebergs, and the increased open water allows icebergs more opportunities to scour the seabed, reducing zoobenthic blue carbon capture and storage. Here the size and variability in benthic blue carbon in mega and macrobenthos was assessed in time and space at Ryder and Marguerite bays of the West Antarctic Peninsula (WAP). In particular the influence of the duration of primary productivity and ice scour are investigated from the shallows to typical shelf depths of 500 m. Ice scour frequency dominated influence on benthic blue carbon at 5 m, to comparable with phytoplankton duration by 25 m depth. At 500 m only phytoplankton duration was significant and influential. WAP zoobenthos was calculated to generate ~10⁷, 4.5 × 10⁶ and 1.6 × 10⁶ tonnes per year (between 2002 and 2015) in terms of production, immobilization and sequestration of carbon respectively. Thus about 1% of annual primary productivity has sequestration potential at the end of the trophic cascade. Polar zoobenthic blue carbon capture and storage responses to sea ice losses, the largest negative feedback on climate change, has been underestimated despite some offsetting of gain by increased ice scouring with more open water. Equivalent survey of Arctic and sub‐Antarctic shelves, for which new projects have started, should reveal the true extent of this feedback and how much its variability contributes to uncertainty in climate models.     

Particle size selection in individuals from epifaunal versus infaunal populations of the nereidid polychaete Neanthes succinea(Polychaeta: Nereididae)

Neanthes succinea (Frey & Leuckart, 1847) is a common nereidid polychaete of both epifaunal and infaunal estuarine habitats. The gut contents of individuals collected from two epifaunal and two infaunal habitats are compared. Our a priori expectation was that individuals from epifaunal habitats would be classified as macrophagous with guts indicating carnivory and/or macroalgal herbivory, while individuals from infaunal habitats would be classified as microphagous with guts indicating deposit feeding. At all four locations gut contents indicated deposit feeding with little indication of macrophagous feeding. Average particle sizes for mineral grains did not differ between the four collection sites. For the two infaunal locations mean size of the mineral grains in gut contents was significantly smaller than ambient sediments. In addition to mineral grains, guts contained diatoms, dinoflagellates, macrophytic detritus, protozoan tests, and a variety of metazoans. Our study demonstrates that caution is necessary when inferring feeding type from morphology and that population and habitat specific differences in diet can occur within the same species.     

Changing patterns of organic carbon and nitrogen accretion on the middle Ebro floodplain (NE Spain)

Sediment profiles from four sites of the middle Ebro River (NE Spain) were examined to describe total organic carbon (TOC) and total nitrogen (TN) accretion patterns during the last century. Such patterns were related to the potential of floodplain substrates to act as C and N sinks. Two oxbow lakes with different hydroperiods and two patches of riparian forest at different successional stages were selected to represent permanently and intermittently flooded habitats, respectively. Total organic carbon (TOC) and total nitrogen (TN) stocks were estimated, whereas Cs-137 dating and aerial photographs were used to determine historical accretion rates. Our results indicate that TOC and TN accretion patterns have been modified during the last century as a result of human impact on natural river processes at both basin and site scales. Since 1963, sediment deposition at the study sites has diminished, regardless of their hydrological connectivity, indicating a reduction in the potential of the middle Ebro floodplains to act as TOC and TN sinks. Although TOC and TN concentrations are markedly higher in recent sediments, accretion rates between 1927 and 1963 still exceed those of recent years. Despite such recent human-induced changes at basin and habitat scale, the middle Ebro floodplains presented relatively high TOC and TN accretion rates, a fact that should be considered in future basin management plans. Based on those results, suggestions to maximize TOC and TN accretion through ecological restoration are provided. Specific actions include the rehabilitation of fine-grained (<2 mm) sediment delivery, land reclamation or dyke reconfiguration.     

Distribution patterns of benthic microalgal standing stock at McMurdo Sound,Antarctica

Summary During the austral summer of 1975–76 and winter of 1977 benthic and water column chlorophyll a and phaeopigments were measured at several sites along the east and west sides of McMurdo Sound, Antarctica. Estimates of in situ primary productivity were made at some McMurdo Sound locations. Additionally, water column samples were collected at 5 stations in the Ross Sea during January, 1976. Standing stock data are analyzed to identify seasonal and spatial patterns. Variability in algal standing stock was related to ambient light levels and appeared to be mediated by ice and snow cover whereby the highest algal standing stock was present under high light conditions (low ice and snow cover, shallow water, summer). Differences in published benthic invertebrate densities appear to be closely allied to differences in benthic primary production, and less so to in situ planktonic ice microalgal production.     

Negative feedback in the cold: ice retreat produces new carbon sinks in Antarctica

Feedbacks on climate change so far identified are predominantly positive, enhancing the rate of change. Loss of sea‐ice, increase in desert areas, water vapour increase, loss of tropical rain forest and the restriction of significant areas of marine productivity to higher latitude (thus smaller geographical zones) all lead to an enhancement of the rate of change. The other major feedback identified, changes in cloud radiation, will produce either a positive feedback, if high level clouds are produced, or a negative feedback if low level clouds are produced. Few significant negative feedbacks have been identified, let alone quantified. Here, we show that the loss of ice shelves and retreat of coastal glaciers around the Antarctic Peninsula in the last 50 years has exposed at least 2.4 × 10⁴ km² of new open water. We estimate that these new areas of open water have allowed new phytoplankton blooms containing a total standing stock of ～5.0 × 10⁵ tonnes of carbon to be produced. New marine zooplankton and seabed communities have also been produced, which we estimate contain ～4.1 × 10⁵ tonnes of carbon. This previously unquantified carbon sink acts as a negative feedback to climate change. New annual productivity, as opposed to standing stock, amounts to 3.5 × 10⁶ tonnes yr^?1 of carbon, of which 6.9 × 10⁵ tonnes yr^?1 deposits to the seabed. By comparison the total aboveground biomasses of lowland American tropical rainforest is 160–435 tonnes ha^?1. Around 50% of this is carbon. On this basis the carbon held in new biomass described here is roughly equivalent to 6000–17 000 ha of tropical rainforest. As ice loss increases in polar regions this feedback will become stronger, and eventually, over thousands to hundreds of thousands of years, over 50 Mtonnes of new carbon could be fixed annually in new coastal phytoplankton blooms and over 10 Mtonnes yr^?1 locked in biological standing stock around Antarctica.     

Predicting the cover and richness of intertidal macroalgae in remote areas: a case study in the Antarctic Peninsula

Antarctica is an iconic region for scientific explorations as it is remote and a critical component of the global climate system. Recent climate change causes a dramatic retreat of ice in Antarctica with associated impacts to its coastal ecosystem. These anthropogenic impacts have a potential to increase habitat availability for Antarctic intertidal assemblages. Assessing the extent and ecological consequences of these changes requires us to develop accurate biotic baselines and quantitative predictive tools. In this study, we demonstrated that satellite‐based remote sensing, when used jointly with in situ ground‐truthing and machine learning algorithms, provides a powerful tool to predict the cover and richness of intertidal macroalgae. The salient finding was that the Sentinel‐based remote sensing described a significant proportion of variability in the cover and richness of Antarctic macroalgae. The highest performing models were for macroalgal richness and the cover of green algae as opposed to the model of brown and red algal cover. When expanding the geographical range of the ground‐truthing, even involving only a few sample points, it becomes possible to potentially map other Antarctic intertidal macroalgal habitats and monitor their dynamics. This is a significant milestone as logistical constraints are an integral part of the Antarctic expeditions. The method has also a potential in other remote coastal areas where extensive in situ mapping is not feasible.     

Preliminary investigation of the contribution of fast-ice algae to the spring phytoplankton bloom in Ellis Fjord,eastern Antarctica

 Algae released from fast-ice in Ellis Fjord, eastern Antarctica, made little contribution to subsequent phytoplankton growth. Dominant taxa in the interior ice community included Nitzschia cylindrus (Grun) Hasle, Navicula glaciei V.H. and a dinoflagellate cyst. Diatom mortality within the ice was high. The algal contribution to the phytoplankton from the fast ice was estimated by calculating the difference between algal biomass in ice cores taken on 14 November with those taken on 18 December 1992. The biomass of sedimenting phytoplankton was estimated using sediment traps; weekly cell counts of water were used to monitor net phytoplankton growth. The low contribution from the fast-ice of Ellis Fjord to the phytoplankton is similar to results from other Antarctic fast-ice communities but is not necessarily reflective of processes occurring within either Antarctic or Arctic pack ice communities. An algal mat growing on the base of the fast-ice had a carbon standing crop of between 0.231 gC m^-2 and 0.022 gC m^-2. Much of this was delivered to the water column as the ice melted while the remainder was exported. Received: 15 March 1995/Accepted: 4 September 1995     

Climate change,glacier retreat and a new ice‐free island offer new insights on Antarctic benthic responses

The Antarctic Peninsula is among the places on Earth that registered major warming in the last 60 yr. Massive ice losses, represented by glacier retreat, ice‐shelf collapses and sea‐ice reduction are among the main impacts of this regional warming. The loss of sea‐bed ice coverage, on the one hand has been affecting benthic assemblages, but on the other it is opening up new areas for benthic colonisation. Potter Cove (South Shetland Islands) offered the opportunity of assessing both processes. We recently reported a sudden shift of benthic assemblages related to increased sedimentation rates caused by glacier retreat. This glacier retreat also uncovered a new island that presents a natural experiment to study Antarctic benthic colonisation and succession. We sampled the new island by photo‐transects taken up to 30 m depth. Here, we report an unexpected benthic assemblage characterised by high species richness, diversity and structural complexity with a well‐developed three‐dimensional structure and epibiotic relationships. Filter feeders comprised the largest trophic group at all depths, mainly ascidians, sponges and bryozoans. Densities were also surprising, recording only six ascidian species with a mean of ～310 ind. m^–2. These values are at least an order of magnitude higher than previous Antarctic reports on early colonisation. This finding challenges the extended idea of a slow and continuous recruitment in Antarctica. However, it also opens the question of whether these complex assemblages could have been present under the glacier in ice‐free refuges that are now exposed to open sea conditions. Under the current scenario of climate change, these results acquire high relevance as they suggest a two‐fold effect of the Antarctic Peninsula warming: the environmental shifts that threaten coastal ecosystems, and also the opening up of new areas for colonisation that may occur at a previously unimagined speed.     

A large carbon pool and small sink in boreal Holocene lake sediments

Model‐based estimates suggest that lake sediments may be a significant, long‐term sink for organic carbon (C) at regional to global scales. These models have used various approaches to predict sediment storage at broad scales from very limited data sets. Here, we report a large‐scale direct assessment of the standing stock and sedimentation rate of C for a representative set of lakes in Finland. The 122 lakes were selected from the statistically selected Nordic Lake Survey database, they cover the entire country and the water quality represents the average lake water quality in Finland. Unlike all prior estimates, these data use sediment cores that comprise the entire sediment record. The data show that within Finland, aquatic ecosystems contain the second largest areal C stocks (19 kg C m^?2) after peatlands (72 kg C m^?2), and exceed by significant amounts stocks in the forest soil (uppermost 75cm; 7.2 kg C m^?2) and woody biomass (3.4 kg C m^?2). Kauppi et al. (1997). The Finnish estimate extrapolated over the boreal region gives a total C pool in lakes 19–27 Pg C, significantly lower than the previous model‐based estimates.     

Modern conophyton‐like microbial mats discovered in Lake Vanda,Antarctica

Lake Vanda is a cold nonturbulent, perennially ice‐covered lake in the valleys of southern Victoria Land, Antarctica. Observations made and samples collected under the 3.5 m ice in 1980 by SCUBA divers reveal that an extensive benthic microbial mat dominated by the filamentous blue‐green algae (cyanobacteria) Phormidium frigidum and Lyngbya martensiana is growing there. As is the case in other Antarctic lakes investigated by us thus far, the mat in Lake Vanda traps and binds sediment and precipitates calcite and is undisturbed by grazers and burrowers. Therefore, stromatolitic laminae are being generated. Unlike the other Antarctic lakes investigated in this region, Lake Vanda has (a) an ice cover and water that transmits significantly more light; (b) an ice cover that is permeable to gases and aeolian sediment; (c) no zone of lift‐off mat where photosynthetically generated oxygen would render the mat buoyant and cause it to separate from the substrate and float away; and (d) mat that has a distinctive pinnacle macrostructure. Although the laminae being laid down by the Lake Vanda mat do not retain the cone and ridge morphology of the living mat, the pinnacle macrostructure of the mat is similar to the Precambrian Conophyton stromatolites as well as microbial structures forming in Yellowstone hot springs, freshwater marshes in the Bahamas, and hypersaline intertidal mats in Baja California, Mexico, and Shark Bay, Australia. This suggests (a) Conophyton‐like structures similar to those abundant during the Precambrian can form under widely varying environmental conditions and (b) high latitudes should not be overlooked as sites of formation of ancient stromatolites.     

Perennially ice-covered Lake Hoare,Antarctica: physical environment,biology and sedimentation

Lake Hoare (77° 38 S, 162° 53 E) is a perennially ice-covered lake at the eastern end of Taylor Valley in southern Victoria Land, Antarctica. The environment of this lake is controlled by the relatively thick ice cover (3–5 m) which eliminates wind generated currents, restricts gas exchange and sediment deposition, and reduces light penetration. The ice cover is in turn largely controlled by the extreme seasonality of Antarctica and local climate. Lake Hoare and other dry valley lakes may be sensitive indicators of short term (< 100 yr) climatic and/or anthropogenic changes in the dry valleys since the onset of intensive exploration over 30 years ago. The time constants for turnover of the water column and lake ice are 50 and 10 years, respectively. The turnover time for atmospheric gases in the lake is 30–60 years. Therefore, the lake environment responds to changes on a 10–100 year timescale. Because the ice cover has a controlling influence on the lake (e.g. light penetration, gas content of water, and sediment deposition), it is probable that small changes in ice ablation, sediment loading on the ice cover, or glacial meltwater (or groundwater) inflow will affect ice cover dynamics and will have a major impact on the lake environment and biota.     

Carbon burial by shallow lakes on the Yangtze floodplain and its relevance to regional carbon sequestration

Floodplain lakes may play an important role in the cycling of organic matter at the landscape scale. For those lakes on the middle and lower reaches of the Yangtze (MLY) floodplain which are subjected to intense anthropogenic disturbance, carbon burial rates should, theoretically, be substantial due to the high nutrient input, increased primary production and high sediment accumulation rates. There are more than 600 lakes >1 km² on the Yangtze floodplain including 18 lakes >100 km² and most are shallow and eutrophic. ²¹⁰Pb‐dated cores were combined with total organic carbon (TOC) analyses to determine annual C accumulation rates (C AR; g C m^?2 yr^?1) and the total C stock (since ~1850). The sediment TOC content is relatively low with an average <2% in most lakes. C AR ranged from ~5 to 373 g C m^?2 yr^?1, resulting in C standing stocks of 0.60–15.3 kg C m^?2 (mean: ~5 kg C m^?2) since ~1850. A multicore study of Chaohu lake (770 km²) indicated that spatial variability of C burial was not a significant problem for regional upscaling. The possible effect of changes in lake size and catchment land use on C burial was examined at Taibai lake and indicated that lake shrinkage and declining arable agriculture had limited effects on C AR. The organic C standing stock in individual lakes is, however, significantly dependent on lake size, allowing a simple linear scaling for all the MLY lakes. Total regional C sequestration was ~80 Tg C since ~1850, equivalent to ~11% of C sequestration by soils, but in ~3% of the land area. Shallow lakes from MLY are a substantial regional C sink, although strong mineralization occurs due to their shallow nature and their role as C sinks is threatened due to lake drainage.     

Control of “blue carbon” storage by mangrove ageing: Evidence from a 66‐year chronosequence in French Guiana

The role of mangroves in the blue carbon stock is critical and requires special focus. Mangroves are carbon‐rich forests that are not in steady‐state equilibrium at the decadal time scale. Over the last decades, the structure and zonation of mangroves have been largely disturbed by coastal changes and land use conversions. The amount of time since the last disturbance is a key parameter determining forest structure, but it has so far been overlooked in mangrove carbon stock projections. In particular, the carbon sequestration rates among mangrove successional ages after (re)establishment are poorly quantified and not used in large‐scale estimations of the blue carbon stock. Here, it is hypothesized that ecosystem age structure significantly modulates mangrove carbon stocks. We analysed a 66‐year chronosequence of the aboveground and belowground biomass and soil carbon stock of mangroves in French Guiana, and we found that in the year after forest establishment on newly formed mud banks, the aboveground, belowground and soil carbon stocks averaged 23.56 ± 7.71, 13.04 ± 3.37 and 84.26 ± 64.14 (to a depth of 1 m) Mg C/ha, respectively. The mean annual increment (MAI) in the aboveground and belowground reservoirs was 23.56 × Age^−0.52 and 13.20 × Age^−0.64 Mg C ha⁻¹ year⁻¹, respectively, and the MAI in the soil carbon reservoir was 3.00 ± 1.80 Mg C ha⁻¹ year⁻¹. Our results show that the plant carbon sink capacity declines with ecosystem age, while the soil carbon sequestration rate remains constant over many years. We suggest that global projections of the above‐ and belowground reservoirs of the carbon stock need to account for mangrove age structures, which result from historical changes in coastal morphology. Our work anticipates joint international efforts to globally quantify the multidecadal mangrove carbon balance based on the combined use of age‐based parametric equations and time series of mangrove age maps at regional scales.     

Stocks and losses of soil organic carbon from Chinese vegetated coastal habitats

Global vegetated coastal habitats (VCHs) represent a large sink for organic carbon (OC) stored within their soils. The regional patterns and causes of spatial variation, however, remain uncertain. The sparsity and regional bias of studies on soil OC stocks from Chinese VCHs have limited the reliable estimation of their capacity as regional and global OC sinks. Here, we use field and published data from 262 sampled soil cores and 181 surface soils to report estimates of soil OC stocks, burial rates and losses of VCHs in China. We find that Chinese mangrove, salt marsh and seagrass habitats have relatively low OC stocks, storing 6.3 ± 0.6, 7.5 ± 0.6, and 1.6 ± 0.6 Tg C (±95% confidence interval) in the top meter of the soil profile with burial rates of 44 ± 17, 159 ± 57, and 6 ± 45 Gg C/year, respectively. The variability in the soil OC stocks is linked to biogeographic factors but is mostly impacted by sedimentary processes and anthropic activities. All habitats have experienced significant losses, resulting in estimated emissions of 94.2–395.4 Tg CO₂e (carbon dioxide equivalent) over the past 70 years. Reversing this trend through conservation and restoration measures has, therefore, great potential in contributing to the mitigation of climate change while providing additional benefits. This assessment, on a national scale from highly sedimentary environments under intensive anthropogenic pressures, provides important insights into blue carbon sink mechanism and sequestration capacities, thus contributing to the synchronous progression of global blue carbon management.     

Planktonic microbial assemblages and the potential effects of metazooplankton predation on the food web of lakes from the maritime Antarctica and sub-Antarctic islands

Antarctica is the continent with the harshest climate on the Earth. Antarctic lakes, however, usually presents liquid water, at least during part of the year or below the ice cover, especially those from the sub-Antarctic islands and the maritime Antarctic region where climatic conditions are less extreme. Planktonic communities in these lakes are mostly dominated by microorganisms, including bacteria and phototrophic and heterotrophic protists, and by metazooplankton, usually represented by rotifers and calanoid copepods, the latter mainly from the genus Boeckella. Here I report and discuss on studies performed during the last decade that show that there is a potential for top–down control of the structure of the planktonic microbial food web in sub-Antarctic and maritime Antarctic lakes. In some of the studied lakes, the effect of copepod grazing on protozoa, either ciliates or flagellates, depending on size of both the predator and the prey, could promote cascade effects that would be transmitted to the bacterioplankton assemblage.