Deniz Özkundakci

Nitrogen and Phosphorus Limitation of Phytoplankton Growth in New Zealand Lakes: Implications for Eutrophication Control

We examine macronutrient limitation in New Zealand (NZ) lakes where, contrary to the phosphorus (P) only control paradigm, nitrogen (N) control is widely adopted to alleviate eutrophication. A review of published results of nutrient enrichment experiments showed that N more frequently limited lake productivity than P; however, stoichiometric analysis of a sample of 121 NZ lakes indicates that the majority (52.9%) of lakes have a mean ratio of total nitrogen (TN) to total phosphorus (TP) (by mass) indicative of potential P-limitation (>15:1), whereas only 14.0% of lakes have mean TN:TP indicative of potential N-limitation (<7:1). Comparison of TN, TP, and chlorophyll a data between 121 NZ lakes and 689 lakes in 15 European Union (EU) countries suggests that at the national scale, N has a greater role in determining lake productivity in NZ than in the EU. TN:TP is significantly lower in NZ lakes across all trophic states, a difference that is driven primarily by significantly lower in-lake TN concentrations at low trophic states and significantly higher TP concentrations at higher trophic states. The form of the TN:TP relationship differs between NZ and the EU countries, suggesting that lake nutrient sources and/or loss mechanisms differ between the two regions. Dual control of N and P should be the status quo for lacustrine eutrophication control in New Zealand and more effort is needed to reduce P inputs.

Relationships between land use and nitrogen and phosphorus in New Zealand lakes

Developing policies to address lake eutrophication requires an understanding of the relative contribution of different nutrient sources and of how lake and catchment characteristics interact to mediate the source–receptor pathway. We analysed total nitrogen (TN) and total phosphorus (TP) data for 101 New Zealand lakes and related these to land use and edaphic sources of phosphorus (P). We then analysed a sub-sample of lakes in agricultural catchments to investigate how lake and catchment variables influence the relationship between land use and in-lake nutrients. Following correction for the effect of co-variation amongst predictor variables, high producing grassland (intensive pasture) was the best predictor of TN and TP, accounting for 38.6% and 41.0% of variation, respectively. Exotic forestry and urban area accounted for a further 18.8% and 3.6% of variation in TP and TN, respectively. Soil P (representing naturally-occurring edaphic P) was negatively correlated with TP, owing to the confounding effect of pastoral land use. Lake and catchment morphology (zmax and lake : catchment area) and catchment connectivity (lake order) mediated the relationship between intensive pasture and in-lake nutrients. Mitigating eutrophication in New Zealand lakes requires action to reduce nutrient export from intensive pasture and quantifying P export from plantation forestry requires further consideration.

Evaluating early-warning indicators of critical transitions in natural aquatic ecosystems

Significance Early-warning indicators (EWIs), statistical metrics of system resilience, have been hypothesized to provide advance warning of sudden shifts in ecosystems, or so-called “regime shifts.” Here we tested this hypothesis for four commonly used EWIs. We used empirical time series from five freshwater ecosystems with documented sudden, persistent transitions hypothesized to represent critical transitions. EWIs were detected in several of these long-term records, and in some cases several years before the transition; however, these EWIs varied in reliability, and agreement between indicators was low. Moreover, their applicability was strongly limited by the requirement for ecosystem-specific knowledge of transition-generating mechanisms and their drivers to choose relevant state variables for analysis. Ecosystems can show sudden and persistent changes in state despite only incremental changes in drivers. Such critical transitions are difficult to predict, because the state of the system often shows little change before the transition. Early-warning indicators (EWIs) are hypothesized to signal the loss of system resilience and have been shown to precede critical transitions in theoretical models, paleo-climate time series, and in laboratory as well as whole lake experiments. The generalizability of EWIs for detecting critical transitions in empirical time series of natural aquatic ecosystems remains largely untested, however. Here we assessed four commonly used EWIs on long-term datasets of five freshwater ecosystems that have experienced sudden, persistent transitions and for which the relevant ecological mechanisms and drivers are well understood. These case studies were categorized by three mechanisms that can generate critical transitions between alternative states: competition, trophic cascade, and intraguild predation. Although EWIs could be detected in most of the case studies, agreement among the four indicators was low. In some cases, EWIs were detected considerably ahead of the transition. Nonetheless, our results show that at present, EWIs do not provide reliable and consistent signals of impending critical transitions despite using some of the best routinely monitored freshwater ecosystems. Our analysis strongly suggests that a priori knowledge of the underlying mechanisms driving ecosystem transitions is necessary to identify relevant state variables for successfully monitoring EWIs.

Latitudinal variation in nutrient stoichiometry and chlorophyll-nutrient relationships in lakes: A global study

We present analysis of variations in relationships between nitrogen (N), phosphorus (P) and chlorophylla (chl-a) in lakes along a gradient of latitude inclusive of tropical, temperate and polar regions. Total nitrogen (TN), total phosphorus (TP), chl-a, latitude and depth data were collated for 1316 lakes situated between 70 ° S and 83 ° N. Latitudinal variation was then analysed for three empirical measures of phytoplankton nutrient limitation and/ or nutrient assimilation. Lastly, chl-a near-maxima conditional on TN and TP abundance were empirically defined for this global dataset using quantile regression. Mean TN:TP increases with distance from the equator. This relationship is independent of variation in either lake depth or trophic state, reflecting latitudinal variation in nutrient cycling processes and/or nutrient sources. There is a negative linear relationship between latitude and chl-a:TN which similarly suggests that N is less abundant relative to phytoplankton growth requirements at lower latitudes. Relative to temperate lakes, the statistical capability of TN and TP to predict chl-a is poor for both tropical and polar lakes, reflecting latitudinal variation in lake ecosystem functioning and the subsequent potential unsuitability of applying relationships derived for temperate lakes elsewhere. Chl-a near-maxima correspond to chl-a:TN and chl-a:TP yields of 0.046:1 and 0.87:1 respectively, although some observations greatly exceed near-maxima, suggesting possible physiologically plastic phytoplankton responses in these exceptional cases. Deficiencies in understanding the mechanisms that drive variation in macro-nutrient stoichiometry and phytoplankton biomass-nutrient relationships across large spatial scales necessitates further landscape-scale research on this topic, particularly in the tropics.

Modelling the response of a highly eutrophic lake to reductions in external and internal nutrient loading

The reduction of macronutrients to levels that limit primary production is often a critical element of mitigating eutrophication and reducing the potential for algal blooms. Lake Okaro has remained highly eutrophic despite an intensive catchment and in-lake restoration programme, including implementation of a constructed wetland, riparian protection, an alum application and application of a modified zeolite mineral (Z2G1) to reduce internal nutrient loading. A one-dimensional process-based ecosystem model (DYRESM-CAEDYM) was used in this study to investigate the need for further nutrient loading reductions of both nitrogen (N) and phosphorus (P). The model was calibrated against field data for a 2-year period and validated over two separate 1-year periods. Model simulations suggest that the trophic status of the lake, measured quantitatively with the Trophic Level Index (TLI), could shift from highly eutrophic to mesotrophic with external and internal loads of both N and P reduced by 75–90%. The magnitude of the nutrient load reductions is indicative of a major challenge in being able to effect transitions across trophic state categories for eutrophic lakes.

Winter severity determines functional trait composition of phytoplankton in seasonally ice‐covered lakes

How climate change will affect the community dynamics and functionality of lake ecosystems during winter is still little understood. This is also true for phytoplankton in seasonally ice-covered temperate lakes which are particularly vulnerable to the presence or absence of ice. We examined changes in pelagic phytoplankton winter community structure in a north temperate lake (Müggelsee, Germany), covering 18 winters between 1995 and 2013. We tested how phytoplankton taxa composition varied along a winter-severity gradient and to what extent winter severity shaped the functional trait composition of overwintering phytoplankton communities using multivariate statistical analyses and a functional trait-based approach. We hypothesized that overwintering phytoplankton communities are dominated by taxa with trait combinations corresponding to the prevailing winter water column conditions, using ice thickness measurements as a winter-severity indicator. Winter severity had little effect on univariate diversity indicators (taxon richness and evenness), but a strong relationship was found between the phytoplankton community structure and winter severity when taxon trait identity was taken into account. Species responses to winter severity were mediated by the key functional traits: motility, nutritional mode, and the ability to form resting stages. Accordingly, one or the other of two functional groups dominated the phytoplankton biomass during mild winters (i.e., thin or no ice cover; phototrophic taxa) or severe winters (i.e., thick ice cover; exclusively motile taxa). Based on predicted milder winters for temperate regions and a reduction in ice-cover durations, phytoplankton communities during winter can be expected to comprise taxa that have a relative advantage when the water column is well mixed (i.e., need not be motile) and light is less limiting (i.e., need not be mixotrophic). A potential implication of this result is that winter severity promotes different communities at the vernal equinox, which may have different nutritional quality for the next trophic level and ecosystem-scale effects.

Phosphorus dynamics in sediments of a eutrophic lake derived from 31P nuclear magnetic resonance spectroscopy

The determination of organic phosphorus (P) compounds in lake sediments can provide information on the potential for internal P loading. Settling seston and vertical sediment core samples from highly eutrophic Lake Okaro, New Zealand, were collected during a mixed winter and stratified summer period, representing, respectively, when the water column was well oxygenated and when the bottom waters were anoxic. Samples were analysed with 31P nuclear magnetic resonance (31P NMR) spectroscopy, which showed that both bottom sediments and settling seston contained orthophosphate, orthophosphate monoesters and diesters, pyrophosphates, polyphosphates and phosphonates. Phosphorus concentration in settling seston increased ~2.5-fold in winter as a result of seasonally induced changes in phytoplankton biomass, with a marked increase in the concentration of orthophosphate. Several potentially bioavailable P compounds in the bottom sediments were identified that were likely to contribute to recycling of P from the sediment to the water column. An ‘apparent half-life’ was used to quantify the time scales on which these compounds were recycled to the overlying water column. Orthophosphate monoesters that include inositol phosphates were the most persistent P compound. On the basis of half-lives, high internal P loadings may persist for more than 20 years, potentially hindering restoration efforts in Lake Okaro.

Parameterisation of sediment geochemistry for simulating water quality responses to long-term catchment and climate changes in polymictic, eutrophic Lake Rotorua, New Zealand

Numerical models of aquatic ecosystems that couple physics and biogeochemistry are valuable tools in aquatic ecosystem research. These models provide opportunities to test theories and to inform environmental management. In this study, we used the dynamic, process-based hydrodynamic-ecological model DYRESM-CAEDYM to simulate key ecosystem processes of Lake Rotorua, New Zealand, for six 8-year periods between 1920 and 2100 in order to evaluate the potential effects of future changes in land use and climate. Longterm variations in external boundary conditions (e.g. inflows) to the lake ecosystem are incorporated by varying the relevant input files in the DYRESMCAEDYM model. However, quantification of internal lake processes, specifically those at the sediment-water interface, presents a major challenge for long-term simulations. The sediment model within CAEDYM is ‘static’, with assumed constant sediment composition and a relatively simplistic process representation for nutrient and oxygen fluxes between sediment and water. Specifically, the model regulates sediment phosphate and ammonium release according to concentrations of oxidising species (i.e. oxygen and nitrate), and temperature in the overlying water layer. Sediment oxygen demand is controlled by dissolved oxygen concentrations and temperature in the water layer overlying the sediments. We used a ‘trial and error’ approach to estimate parameters for calibrating and validating the model, and regression modelling to infer the www.witpress.com, ISSN 1743-3541 (on-line) WIT Transactions on Ecology and The Environment, Vol 164,

Characteristics of consumer trophic resources for Waikato shallow lake food webs

ABSTRACT We characterised the quantity (mass per unit area or volume), quality (cabon:nitrogen ratio) and enrichment (δ15N) of basal food resources and benthic macroinvertebrates in six shallow lakes in the Waikato region, New Zealand, to elucidate patterns related to lake type (peat, riverine), habitat (littoral, mid-lake) and enrichment status (Trophic Level Index >6 or ≤5). Total seston dry mass was dominated by fine material that was correlated with chlorophyll a concentration. Seston was a higher quality food resource than benthic particulate organic matter which had higher biomass overall in littoral than in mid-lake habitats. Biomass of Chironomidae was highest in the less-enriched peat lakes, possibly due to clearer water supporting benthic algae in shallow littoral areas. Our results highlight the potential role of multiple bottom-up influences on shallow lake food webs. The more abundant and higher quality pelagic food resources in highly enriched lakes may have implications for resilience to non-indigenous fish invasion.

Patterns and drivers of spatio-temporal variability of suspended sediment in the Waikato lakes, New Zealand

ABSTRACT Quantifying and interpreting variability in ecological systems allows us to better understand the driving forces that determine ecosystems functioning. We utilised earth observation-based remote sensing to determine total suspended sediment (TSS) concentrations for 109 lakes over 16 years. The coefficient of variation (CV) was used to quantify variability of TSS at three temporal scales (monthly, annual and long-term) and at the horizontal spatial scale. CV values varied between 0.14 and 1.09. Drivers of observed variability patterns were explored using five lake morphological and catchment characteristics. Lake area was the most important variable to predict variability of TSS. De-vegetated lakes displayed significantly higher TSS concentrations than vegetated lakes; however, aquatic vegetation was found to moderate only spatial variability of TSS. As spatio-temporal variability can be assumed to drive key ecosystem functions, determining characteristic variability patterns for various attributes should be a prerequisite when setting management goals and developing monitoring regimes.