Microbial food web dynamics were determined during the onset of several spring phytoplankton blooms induced by natural iron fertilization off Kerguelen Island in the Southern Ocean (KEOPS2). The ...abundances of heterotrophic bacteria and heterotrophic nanoflagellates, bacterial heterotrophic production, bacterial respiration, and bacterial growth efficiency, were consistently higher in surface waters of the iron-fertilized sites than at the reference site in HNLC (high nutrient low chlorophyll) waters. The abundance of virus-like particles remained unchanged, but viral production increased by a factor of 6 in iron-fertilized waters. Bacterial heterotrophic production was significantly related to heterotrophic nanoflagellate abundance and viral production across all sites, with bacterial production explaining about 70 and 85%, respectively, of the variance of each in the mixed layer (ML). Estimated rates of grazing and viral lysis, however, indicated that heterotrophic nanoflagellates accounted for a substantially higher loss of bacterial production (50%) than viruses (11%). Combining these results with rates of primary production and export determined for the study area, a budget for the flow of carbon through the microbial food web and higher trophic levels during the early (KEOPS2) and the late phase (KEOPS1) of the Kerguelen bloom is provided.
The corallivorous Crown-of-Thorns Starfish (CoTS, Acanthaster spp.) has been linked with the widespread loss of scleractinian coral cover on Indo-Pacific reefs during periodic population outbreaks. ...Here, we re-examine CoTS consumption by coral reef fish species by using new DNA technologies to detect Pacific Crown-of-Thorns Starfish (Acanthaster cf. solaris) in fish faecal and gut content samples. CoTS DNA was detected in samples from 18 different coral reef fish species collected on reefs at various stages of CoTS outbreaks in the Great Barrier Reef Marine Park, nine of which had not been previously reported to feed on CoTS. A comprehensive set of negative and positive control samples confirmed that our collection, processing and analysis procedures were robust, although food web transfer of CoTS DNA cannot be ruled out for some fish species. Our results, combined with the (i) presence of CoTS spines in some samples, (ii) reported predation on CoTS gametes, larvae and settled individuals, and (iii) known diet information for fish species examined, strongly indicate that direct fish predation on CoTS may well be more common than is currently appreciated. We provide recommendations for specific management approaches to enhance predation on CoTS by coral reef fishes, and to support the mitigation of CoTS outbreaks and reverse declines in hard coral cover.
This paper presents results on the spatial and temporal distribution patterns of mesozooplankton in the naturally fertilized region to the east of the Kerguelen Islands (Southern Ocean) visited at ...early bloom stage during the KEOPS2 survey (15 October to 20 November 2011). The aim of this study was to compare the zooplankton response in contrasted environments localized over the Kerguelen Plateau in waters of the east shelf and shelf edge and in productive oceanic deep waters characterized by conditions of complex circulation and rapidly changing phytoplankton biomass. The mesozooplankton community responded to the spring bloom earlier on the plateau than in the oceanic waters, where complex mesoscale circulation stimulated initial more or less ephemeral blooms before a broader bloom extension. Taxonomic compositions showed a high degree of similarity across the whole region, and the populations initially responded to spring bloom with a large production of larval forms increasing abundances, without biomass changes. Taxonomic composition and stable isotope ratios of size-fractionated zooplankton indicated the strong domination of herbivores, and the total zooplankton biomass values over the survey presented a significant correlation with the integrated chlorophyll concentrations in the mixed layer. The biomass stocks observed at the beginning of the KEOPS2 cruise were around 1.7 g C m−2 above the plateau and 1.2 g C m−2 in oceanic waters. Zooplankton biomass in oceanic waters remained on average below 2 g C m−2 over the study period, except for one station in the Polar Front zone (F-L), whereas zooplankton biomasses were around 4 g C m−2 on the plateau at the end of the survey. The most remarkable feature during the sampling period was the stronger increase in abundance in the oceanic waters (25 × 103 to 160 × 103 ind m−2) than on the plateau (25 × 103 to 90 × 103 ind m−2). The size structure and taxonomic distribution patterns revealed a cumulative contribution of various larval stages of dominant copepods and euphausiids particularly in the oceanic waters, with clearly identifiable stages of progress during a Lagrangian time series survey. The reproduction and early stage development of dominant species were sustained by mesoscale-related initial ephemeral blooms in oceanic waters, but growth was still food-limited and zooplankton biomass stagnated. In contrast, zooplankton abundance and biomass on the shelf were both in a growing phase, at slightly different rates, due to growth under sub-optimal conditions. Combined with our observations during the KEOPS1 survey (January–February 2005), the present results deliver a consistent understanding of patterns in mesozooplankton abundance and biomass from early spring to summer in the poorly documented oceanic region east of the Kerguelen Islands.
Seasonal variation in coral reef macroalgal size and condition is well documented, yet seasonal variability of herbivory on macroalgae by coral reef fishes is unknown. Herbivore feeding intensity was ...quantified monthly on an inner-shelf reef on the Great Barrier Reef, using Sargassum bioassays. Removal rates of transplants displayed high levels of variation with significantly higher rates of removal during the summer months. Differences in Sargassum plant size and condition suggest that the variability in herbivore feeding intensity is attributed primarily to the variation in the condition of the macroalgae, especially epiphyte loads. The dramatic changes in macroalgal removal reveal a considerable decrease in herbivore activity in the winter. This highlights the clear distinction between ‘summer' and ‘winter' months in terms of reef processes, emphasizing the high seasonal variation in macroalgal removal rates at different time of the year.
Although the Southern Ocean is considered a high-nutrient, low-chlorophyll (HNLC) area, massive and recurrent blooms are observed over and downstream of the Kerguelen Plateau. This mosaic of blooms ...is triggered by a higher iron supply resulting from the interaction between the Antarctic Circumpolar Current and the local bathymetry. Net primary production, N uptake (NO3− and NH4+), and nitrification rates were measured at eight stations in austral spring 2011 (October–November) during the KEOPS 2 cruise in the Kerguelen Plateau area. Natural iron fertilization stimulated primary production, with mixed layer integrated net primary production and growth rates much higher in the fertilized areas (up to 315 mmol C m−2 d−1 and up to 0.31 d−1 respectively) compared to the HNLC reference site (12 mmol C m−2 d−1 and 0.06 d−1 respectively). Primary production was mainly sustained by nitrate uptake, with f ratios (corresponding to NO3−-uptake / (NO3−-uptake + NH4+-uptake)) lying at the upper end of the observations for the Southern Ocean (up to 0.9). We report high rates of nitrification (up to ~ 3 μmol N L−1 d−1, with ~ 90 % of them < 1 μmol N L−1 d−1) typically occurring below the euphotic zone, as classically observed in the global ocean. The specificity of the studied area is that at most of the stations, the euphotic layer was shallower than the mixed layer, implying that nitrifiers can efficiently compete with phytoplankton for the ammonium produced by remineralization at low-light intensities. Nitrate produced by nitrification in the mixed layer below the euphotic zone is easily supplied to the euphotic zone waters above, and nitrification sustained 70 ± 30 % of the nitrate uptake in the productive area above the Kerguelen Plateau. This complicates estimations of new production as potentially exportable production. We conclude that high productivity in deep mixing system stimulates the N cycle by increasing both assimilation and regeneration.
We report on the zonal variability of mesopelagic particulate organic carbon remineralization and deep carbon transfer potential during the Kerguelen Ocean and Plateau compared Study 2 expedition ...(KEOPS 2; October-November 2011) in an area of the polar front supporting recurrent massive blooms from natural Fe fertilization. Mesopelagic carbon remineralization (MR) was assessed using the excess, non-lithogenic particulate barium (Baxs) inventories in mesopelagic waters and compared with bacterial production (BP), surface primary production (PP) and export production (EP). Results for this early season study are compared with the results obtained during a previous study (2005; KEOPS 1) for the same area at a later stage of the phytoplankton bloom. Our results reveal the patchiness of the seasonal advancement and of the establishment of remineralization processes between the plateau (A3) and polar front sites during KEOPS 2. For the Kerguelen plateau (A3 site) we observe a similar functioning of the mesopelagic ecosystem during both seasons (spring and summer), with low and rather stable remineralization fluxes in the mesopelagic column (150-400 m). The shallow water column (~500 m), the lateral advection, the zooplankton grazing pressure and the pulsed nature of the particulate organic carbon (POC) transfer at A3 seem to drive the extent of MR processes on the plateau. For deeper stations (>2000 m) located on the margin, inside a polar front meander, as well as in the vicinity of the polar front, east of Kerguelen, remineralization in the upper 400 m in general represents a larger part of surface carbon export. However, when considering the upper 800 m, in some cases, the entire flux of exported carbon is remineralized. In the polar front meander, where successive stations form a time series, two successive events of particle transfer were evidenced by remineralization rates: a first mesopelagic and deep transfer from a past bloom before the cruise, and a second transfer expanding at mesopelagic layers during the cruise. Regarding the deep carbon transfer efficiency, it appeared that above the plateau (A3 site) the mesopelagic remineralization was not a major barrier to the transfer of organic matter to the seafloor (close to 500 m). There, the efficiency of carbon transfer to the bottom waters (>400 m) as assessed by PP, EP and MR fluxes comparisons reached up to 87% of the carbon exported from the upper 150 m. In contrast, at the deeper locations, mesopelagic remineralization clearly limited the transfer of carbon to depths of >400 m. For sites at the margin of the plateau (station E-4W) and the polar front (station F-L), mesopelagic remineralization even exceeded upper 150 m export, resulting in a zero transfer efficiency to depths >800 m. In the polar front meander (time series), the capacity of the meander to transfer carbon to depth >800 m was highly variable (0 to 73%). The highest carbon transfer efficiencies in the meander are furthermore coupled to intense and complete deep (>800 m) remineralization, resulting again in a near-zero, deep (>2000 m) carbon sequestration efficiency there.
Despite the widely accepted importance of fish herbivory on coral reefs, few studies have considered the temporal variability in the nature of algal–herbivore interactions. We therefore quantified ...monthly feeding intensity onSargassumsp. bioassays for 12 mo with remote underwater video cameras deployed to identify the herbivores responsible for macroalgal removal on an inshore island of the Great Barrier Reef, Australia. Significantly higher removal rates were observed during the summer months whereas winter months were characterized by 4 times lower removal rates. However, rather than being simply changes in the feeding activity of a single species, this temporal pattern in herbivory also incorporated changes in the species responsible for the removal ofSargassum. Video analyses revealed that, of the 43 herbivore species recorded from the bay, only 3 played a significant role inSargassumremoval:Kyphosus vaigiensis,Naso unicornisandScarus rivulatus.K. vaigiensis, a rudderfish, was primarily responsible for the removal ofSargassumduring the summer months (83% of the total recorded bites; 85 553 bites). There was almost no feeding activity onSargassumbyK. vaigiensisduring the winter months (82 bites). However, there was a reciprocal increase in feeding intensity onSargassumby parrotfishes in the winter months, particularlyS. rivulatus(71 bites during summer versus 2884 bites in winter). This temporal variability in herbivore functional roles suggests that functional redundancy on reefs may be less than previously assumed in that the feeding activities of fishes may be both spatially and temporally constrained.
We simultaneously measured bacterial production (BP), bacterial respiration (BR), alkaline phosphatase activity (
phos
) and ectoaminopeptidase activity (
prot
) in relation to biogeochemical ...parameters, nutritive resources and in situ temperature over a 1-year survey at the long-term observatory the SOLEMIO station (Marseille bay, NW Mediterranean Sea). Despite its proximity to the coast, oligotrophic conditions prevailed at this station (yearly mean of Chl
a
= 0.43 μg dm
−3
, NO
3
= 0.55 μmol dm
−3
and PO
4
= 0.04 μmol dm
−3
). Episodic meteorological events (dominant winds, inputs from the Rhone River) induced rapid oscillations (within 15 days) in temperature and sometimes salinity that resulted in rapid changes in phytoplankton succession and a high variability in C/P ratios within the particulate and dissolved organic matter. Throughout the year, BP ranged from 0.01 to 0.82 μg C dm−
3
h−
1
and bacterial growth efficiency varied from 1 to 39 %, with higher values in summer. Enrichment experiments showed that BP was limited most of the year by phosphorus availability (except in winter). A significant positive correlation was found between in situ temperature, BP, BR and
phos
. Finally, we found that temperature and phosphate availability were the main factors driving heterotrophic bacterial activity and thus play a fundamental role in carbon fluxes within the marine ecosystem.
The dynamic nature of coral reefs offers a rare opportunity to examine the response of ecosystems to disruption due to climate change. In 1998, the Great Barrier Reef experienced widespread coral ...bleaching and mortality. As a result, cryptobenthic fish assemblages underwent a dramatic phase-shift. Thirteen years, and up to 96 fish generations later, the cryptobenthic fish assemblage has not returned to its pre-bleach configuration. This is despite coral abundances returning to, or exceeding, pre-bleach values. The post-bleach fish assemblage exhibits no evidence of recovery. If these short-lived fish species are a model for their longer-lived counterparts, they suggest that (1) the full effects of the 1998 bleaching event on long-lived fish populations have yet to be seen, (2) it may take decades, or more, before recovery or regeneration of these long-lived species will begin, and (3) fish assemblages may not recover to their previous composition despite the return of corals.
The majority of coral reef goby species are short-lived, with some highly abundant species living less than 100 d. To understand the role and consequences of this extreme life history in shaping ...coral reef fish populations, we quantitatively documented the structure of small reef fish populations over a 26-month period (>14 short-lived fish generations) at an inshore reef on the Great Barrier Reef, Australia. Most species with life spans >1 yr, such as pomacentrids, exhibited a peak in recruitment during the austral summer, driving seasonal changes in the small fish community composition. In contrast, there were no clear changes in goby community composition, despite the abundance of short-lived, high turnover species. Species of
Eviota
, the most abundant gobiid genus observed, showed remarkably similar demographic profiles year-round, with consistent densities of adults as well as recently recruited juveniles. Our results demonstrate ongoing recruitment of these small cryptic fishes, which appears to compensate for an exceptionally short life span on the reef. Our results suggest that gobiid populations are able to overcome demographic limitations, and by maintaining reproduction, larval survival and recruitment throughout the year, they may avoid population bottlenecks. These findings also underline the potential trophodynamic importance of these small species; because of this constant turnover,
Eviota
species and other short-lived fishes may be particularly valuable contributors to the flow of energy on coral reefs, underpinning the year-round trophic structure.