Impacts of internal waves on the twilight zone @ ICRS 2016, Hawaii (Dr Wyatt)

Ecological and biogeochemical impacts of internal waves on mesophotic coral ecosystems: testing eddy correlation and isotope approaches, Iriomote, Japan

Alex S.J. Wyatt1*, Toshihiro Miyajima1, James J. Leichter2, Tohru Naruse3, Tomohiro Kuwae4, Shoji Yamamoto5, Naomi Satoh1, Toshi Nagata1

1Department of Chemical Oceanography, Atmosphere and Ocean Research Institute, The University of Tokyo, Kashiwa, Chiba, JAPAN
2Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California, USA
3Tropical Biosphere Research Center, University of the Ryukyus, Taketomi, Japan
4Coastal and Estuarine Environment Research Group, Port and Airport Research Institute (PARI), Nagase, Yokosuka, JAPAN
5Department of Earth and Planetary Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan

While mesophotic coral ecosystems (MCE) may be protected or damped from disturbances impacting shallower reefs insufficient information is available on the environmental conditions supporting these ‘deep water refugia’. Nutrient inputs and recycling have rarely been quantified over MCE but may differ fundamentally to that of shallow counterparts due to the reduction in light and increasing reliance on oceanic nutrients, leading to increased heterotrophy over autotrophy at species and ecosystem levels and stronger links to oceanic processes. For instance, due to the depth of MCE relative to typical water column density stratification, internal waves may be a highly significant process depending on community aspect and exposure. Preliminary observations of MCE along a continuum of oceanic exposure in Funauki Bay, Iriomote, Japan indicate that ocean-exposed MCE are subject to semi-diurnal temperature oscillations of up to 4 C during summer (range 23 – 29 deg C), while inner MCE occur shallower in more turbid but stable environments. Oceanic exposure along the bay may determine both the distribution and function of spatially extensive, but relatively homogenous, communities dominated by Leptoseris sp. or Acropora ?horrida. Combining bulk and compound-specific stable isotope analyses, depth-specific radioisotope markers such as radiocarbon, and eddy correlation experiments in these habitat promises a useful approach for elucidating the functional importance of internal waves in the development and persistence of MCE at local to regional scales.

Ecosystem inputs and recycling over coral reefs @ 3rd APCRS, 2014 (Dr Wyatt)

Functional understanding of ecosystem-scale inputs and recycling over coral reef communities from stable isotope analyses of organic matter

Alex S.J. Wyatt1*, James J. Leichter2, Benoit Thibodeau1, Toshihiro Miyajima1, Craig A. Carlson3, Craig E. Nelson4, Toshi Nagata1

1Marine Biogeochemistry Laboratory, Department of Chemical Oceanography, Atmosphere & Ocean Research Institute, The University of Tokyo, Kashiwa, Chiba, JAPAN
2Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California, USA
3University of California at Santa Barbara, Santa Barbara, California, USA
4Center for Microbial Oceanography: Research and Education, University of Hawai’i, USA

Stable isotope analyses (SIA) are an increasingly useful tool for understanding functional links between water flow and nutrient cycling over coral reefs, including relative fluxes of oceanic and reef-derived material. SIA have suggested that oceanic particulate organic matter (POM) flowing over reefs can be rapidly metabolized, with the subsequent release of remineralized inorganic nutrients, as well reef-derived POM, representing a significant resource for downstream communities. High oceanic concentrations of dissolved organic matter (DOM) relative to POM suggests DOM may be an even more significant resource, especially around low-POM reefs such as mid-ocean islands and atolls. However, DOM fluxes have rarely been quantified, perhaps due to the refractory nature of oceanic DOM and difficulties linking small concentration changes with spatial changes in both hydrodynamics and macro- and microbial communities. Our Lagrangian studies of DOM around Moorea, French Polynesia and Ishigaki Island, Japan suggest that DOM changes occurring over short spatial scales reflect a balance between uptake and release. SIA further suggest that the release of reef-derived DOM (i.e. enriched in 13C), perhaps relatively labile and from nitrogen fixing organisms (i.e. depleted in 15N), may promote nutrient recycling and supply to downstream communities. Linking SIA and local hydrodynamics offers a promising path towards elucidating the relative functional importance of oceanic and reef-level processes for reef communities.

Key words: dissolved organic matter, fluxes, particulate organic matter, recycling, stable isotope analyses

Formation and maintenance of high-nitrate, low pH layers (Waite et al., Biogeosciences)

Formation and maintenance of high-nitrate, low pH layers in the eastern Indian Ocean and the role of nitrogen fixation

A. M. Waite, V. Rossi, M. Roughan, B. Tilbrook, P. A. Thompson, M. Feng, A. S. J. Wyatt, and E. J. Raes

We investigated the biogeochemistry of low dissolved oxygen high-nitrate (LDOHN) layers forming against the backdrop of several interleaving regional water masses in the eastern Indian Ocean, off northwest Australia adjacent to Ningaloo Reef. These water masses, including the forming Leeuwin Current, have been shown directly to impact the ecological function of Ningaloo Reef and other iconic coastal habitats downstream. Our results indicate that LDOHN layers are formed from multiple subduction events of the Eastern Gyral Current beneath the Leeuwin Current (LC); the LC originates from both the Indonesian Throughflow and tropical Indian Ocean. Density differences of up to 0.025 kg m−3 between the Eastern Gyral Current and the Leeuwin Current produce sharp gradients that can trap high concentrations of particles (measured as low transmission) along the density interfaces. The oxidation of the trapped particulate matter results in local depletion of dissolved oxygen and regeneration of dissolved nitrate (nitrification). We document an associated increase in total dissolved carbon dioxide, which lowers the seawater pH by 0.04 units. Based on isotopic measurements (δ15N and δ18O) of dissolved nitrate, we determine that ~ 40–100% of the nitrate found in LDOHN layers is likely to originate from nitrogen fixation, and that, regionally, the importance of N-fixation in contributing to LDOHN layers is likely to be highest at the surface and offshore.

Coral reef POM dynamics (Wyatt et al., L&O)

Particulate nutrient fluxes over a fringing coral reef: Source-sink dynamics inferred from carbon to nitrogen ratios and stable isotopes

Alex S. J. Wyatt, Ryan J. Lowe, Stuart Humphries and Anya M. Waite

We examined spatial and temporal variations in particulate organic matter (POM) dynamics over a fringing coral reef (Ningaloo Reef) in Western Australia during the austral autumn and spring. Total POM concentrations generally did not differ between seasons or reef zones, but the composition of POM, in terms of carbon isotope ratios (δ13C-POM), carbon to nitrogen ratios (C : N), and fatty acids, changed consistently in water flowing across the reef. Both δ13C-POM and C : N increased from the fore reef to the reef flat and lagoon, −23.0‰ to −20.1‰ and 7.31 to 8.34, respectively. Average rates of net POM uptake by the reef community were highest over the reef crest (4 to 30 mmol N m−2 d−1 and 6 to 130 mmol C m−2 d−1), with a Bayesian isotope model confirming independent measurements of high uptake rates of allochthonous POM (oceanic phyto- and zooplankton). In contrast, over the reef flat, net release of POM was observed (−4 to −5 mmol N m−2 d−1 and −50 mmol C m−2 d−1), with gross release rates (estimated as −6 to −8 mmol N m−2 d−1 and −30 to −90 mmol C m−2 d−1) indicating that the release of autochthonous POM may be of similar magnitude to allochthonous uptake. Examining POM dynamics in terms of gross fluxes reinforces the dependence of coral reef systems on oceanographic processes for allochthonous POM supply, as well as highlighting the potential for autochthonous POM production to supply nutrients to benthic and pelagic communities downstream.

Coral reef nutrient dynamics (Wyatt et al., L&O)

Oceanographic forcing of nutrient uptake and release over a fringing coral reef

Alex S. J. Wyatt, James L. Falter, Ryan J. Lowe, Stuart Humphries and Anya M. Waite

Nitrate and nitrite (NOx) and phosphate (PO4) dynamics over Ningaloo Reef, Western Australia, are shown to depend on oceanographic forcing of coupled mass transfer limited (MTL) gross uptake and gross release from remineralized oceanic particulate organic matter (POM). Estimates of gross release rates increased significantly with increasing POM uptake and were of the same order as gross uptake rates. Gross uptake rates increased significantly with increasing oceanic concentrations and wave energy dissipation, were 35–80% higher over the reef crest (7–9 mmol NOx m−2 d−1 and 4–5 mmol PO4 m−2 d−1), and were significantly correlated with independent estimates of POM-mediated gross NOx uptake, supporting both MTL uptake and the strong role of oceanic POM supply. The relative supply of NOx and POM was linked to the seasonal dynamics of a regional current system. In late spring, upwelling associated with seasonally strong equator-ward winds led to increased NOx concentrations (0.71 ± 0.2 µmol L−1), POM < NOx and the reef was a net nutrient sink (5390 mmol NOxm−1 d−1 and 270 mmol PO4 m−1 d−1). In contrast, during the autumn, NOx was low (0.16 ± 0.06 µmol L−1), but POM > NOx and the reef was a net nutrient source (−7060 mmol NOx m−1 d−1 and −730 mmol PO4 m−1 d−1). The autumn enhancement of oceanic POM supply to the reef can be attributed to a regional phytoplankton bloom associated with acceleration of the oligotrophic Leeuwin Current, which may result in a significant supply of dissolved nutrients to downstream communities.

Picoplankton and virus uptake by a coral reef (Patten et al., Coral Reefs)

Uptake of picophytoplankton, bacterioplankton and virioplankton by a fringing coral reef community (Ningaloo Reef, Australia)

Nicole L. Patten, Alex S.J. Wyatt, Ryan J. Lowe, Anya M. Waite

We examined the importance of picoplankton and virioplankton to reef trophodynamics at Ningaloo Reef, (north-western Australia), in May and November 2008. Picophytoplankton (Prochlorococcus,Synechococcus and picoeukaryotes), bacterioplankton (inclusive of bacteria and Archaea), virioplankton and chlorophyll a (Chl a) were measured at five stations following the consistent wave-driven unidirectional mean flow path of seawater across the reef and into the lagoon.Prochlorococcus, Synechococcus, picoeukaryotes and bacterioplankton were depleted to similar levels (~40% on average) over the fore reef, reef crest and reef flat (=‘active reef’), with negligible uptake occurring over the sandy bottom lagoon. Depletion of virioplankton also occurred but to more variable levels. Highest uptake rates, m, of picoplankton occurred over the reef crest, while uptake coefficients, S (independent of cell concentration), were similarly scaled over the reef zones, indicating no preferential uptake of any one group. Collectively, picophytoplankton, bacterioplankton and virioplankton accounted for the uptake of 29 mmol C m−2 day−1, with Synechococcuscontributing the highest proportion of the removed C. Picoplankton and virioplankton accounted for 1–5 mmol N m−2 day−1 of the removed N, with bacterioplankton estimated to be a highly rich source of N. Results indicate the importance of ocean–reef interactions and the dependence of certain reef organisms on picoplanktonic supply for reef-level biogeochemistry processes.

Keywords: Coral reef, Picoplankton, Virus, Uptake, Ningaloo Reef, Indian Ocean

Coral reef phytoplankton fluxes (Wyatt et al., MEPS)

Particulate nutrient fluxes over a fringing coral reef: relevant scales of phytoplankton production and mechanisms of supply

Alex S. J. Wyatt, Ryan J. Lowe, Stuart Humphries, Anya M. Waite

Seasonal observations of phytoplankton uptake at Ningaloo Reef, Western Australia, reinforce the importance of particulate organic nitrogen (PON) and carbon (POC) in reef nutrient budgets and identify wave action and the dynamics of regional currents (over a range of temporal and spatial scales) as important factors determining plankton supply to the reef. Phytoplankton uptake rates, calculated from declining chlorophyll a concentrations as water moved over the reef, appeared to be near the physical limits of mass transfer. Phytoplankton-derived PON flux of 2 to 5 mmol N m–2 d–1 was on the order of that typical for dissolved N uptake—confirming that particle feeding may supply the N missing in reef N budgets—while POC flux of 14 to 27 mmol C m–2 d–1 was on the order of net community metabolism. Phytoplankton supply was highly variable at daily-to-seasonal time scales in response to the dynamics of a regional current system dominated by the downwelling-favourable Leeuwin Current (LC). Acceleration of the LC in the austral autumn may supply as much phytoplankton to the reef as sporadic upwelling associated with the Ningaloo Current (NC) in summer. The ocean catchment concept is introduced as a basis for examining the spatial scale of pelagic processes influencing benthic systems: every day, Ningaloo may completely consume the phytoplankton over 87 km2 of LC water, compared to only 20 km2 of NC water. Production within this catchment appears insufficient to maintain offshore phytoplankton concentrations, and advection of remotely sourced production into the catchment is required to balance reef uptake. A functional dependence by reef organisms on externally sourced ocean productivity increases the potential scale at which human- or climatically induced changes may affect reef communities and suggests that processes such as changes in offshore currents and plankton communities require further consideration in reef-level biogeochemistry.

KEY WORDS: Ningaloo Reef · Nutrient budget · Oceanographic forcing · Particulate organic carbon · Particulate organic nitrogen · Leeuwin Current · Ocean catchment · Upwelling