The ecosystem of a mesotrophic lake-II. Interactions among nutrient load,river flow,and epilimnion depth

The management variables which primarily affect phytoplankton biomass (as chl-a) in Lake Mjøsa, Norway, are total phosphorus loading (TP) and the timing and volume of water through flow (by active storage reservoirs). The response of the lake to changes in these factors is studied using a simulation model of the lake ecosystem. Chl-a responses from both observed data and the simulated results are extracted by multiple regression. Results show that decreasing TP load decreases chl-a, but less at low TP levels (< 10 mg TP · m^–3). There is also a certain time period for peak river flow which gives the least yield of chl-a per unit TP. This time period occurs in early summer (i.e., around June 10) if the total phosphorus load is low, and later if the load is high. Both observations and simulation results show that a high water flow increases chl-a at low epilimnion depths (< 15 m), but that the same high water flow decreases chl-a when epilmnion is deep.     

Phytoplankton biomass size structure and its regulation in the Southern Yellow Sea (China): Seasonal variability

Phytoplankton size structure plays a significant role in controlling the carbon flux of marine pelagic ecosystems. The mesoscale distribution and seasonal variation of total and size-fractionated phytoplankton biomass in surface waters, as measured by chlorophyll a (Chl a), was studied in the Southern Yellow Sea using data from four cruises during 2006–2007. The distribution of Chl a showed a high degree of spatial and temporal variation in the study area. Chl a concentrations were relatively high in the summer and autumn, with a mean of 1.42 and 1.27 mg m⁻³, respectively. Conversely, in the winter and spring, the average Chl a levels were only 0.98 and 0.99 mg m⁻³. Total Chl a showed a clear decreasing gradient from coastal areas to the open sea in the summer, autumn and winter cruises. Patches of high Chl a were observed in the central part of the Southern Yellow Sea in the spring due to the onset of the phytoplankton bloom. The eutrophic coastal waters contributed at least 68% of the total phytoplankton biomass in the surface layer. Picophytoplankton showed a consistent and absolute dominance in the central region of the Southern Yellow Sea (>40%) in all of the cruises, while the proportion of microphytoplankton was the highest in coastal waters. The relative proportions of pico- and nanophytoplankton decreased with total biomass, whereas the proportion of the micro-fraction increased with total biomass. Relationships between phytoplankton biomass and environmental factors were also analysed. The results showed that the onset of the spring bloom was highly dependent on water column stability. Phytoplankton growth was limited by nutrient availability in the summer due to the strong thermocline. The combined effects of P-limitation and vertical mixing in the autumn restrained the further increase of phytoplankton biomass in the surface layer. The low phytoplankton biomass in winter was caused by vertical dispersion due to intense mixing. Compared with the availability of nutrients, temperature did not seem to cause direct effects on phytoplankton biomass and its size structure. Although interactions of many different environmental factors affected phytoplankton distributions, hydrodynamic conditions seemed to be the dominant factor. Phytoplankton size structure was determined mainly by the size-differential capacity in acquiring resource. Short time scale events, such as the spring bloom and the extension of Yangtze River plume, can have substantial influences, both on the total Chl a concentration and on the size structure of the phytoplankton.     

The short term response to eutrophication abatement

We compare results of a new model for predicting the short term inter annual changes in chlorophyll-a (chl-a) in lakes after reductions in total phosphorus (TP) to predictions made by least squares regression models. In the new method, slopes of chl-a/TP graphs (both axes in mg · m^–3) are depicted in frequency diagrams and used to extract information on the expected, short term chl-a/TP response. The short term response for nine shallow (< 10 m deep) and nutrient rich lakes to changes in TP was found to be: Chl-a = 0.49 · TP + 17.3, and for nine deep, P-limited lakes: Chl-a = 0.08 · TP + 3.5. If the TP-reduction is known to be greater than 10 mg · m^–3, the expected slope increases to 0.58 for shallow lakes and to 0.26 for deep lakes. The slope, 0.58, is 8% lower than the slope for the long term response calculated by regression for the shallow lakes. For deep lakes the slope, 0.26, is 2 to 3 times higher than that calculated by regression, indicating that reductions in TP for deep lakes give greater effects than least squares regression equations suggest. We have also calculated the reduction in TP which will give about 80% probability that a reduction in chl-a will be observed next year. For shallow, P-limited lakes this reduction is about 30 mg · m^–3 (5% of average initial in-lake TP concentration), and for deep lakes about 14 mg · m^–3 (35% of average initial in-lake TP concentration).     

Harmful algal blooms (HABs) in Daya Bay, China: An in situ study of primary production and environmental impacts

A month-long investigation of phytoplankton biomass and primary production (PP) was carried out during a harmful algal bloom (HAB) in Daya Bay, China, in 2003. During the bloom, the phytoplankton community was dominated by Scrippsiella trochoidea and Chattonella marina. The phytoplankton biomass (Chl a) and PP reached peak levels of 519.21 mg m⁻³ and 734.0 mgC m⁻³ h⁻¹, respectively. Micro-phytoplankton was the key contributor to Chl a and PP in a cage-culture area and in the adjacent HAB-affected waters, with percentages of up to 82.91% and 84.94%, respectively. The HAB had complicated relationships with hydrological and meteorological factors in Daya Bay. However, the water around the cage-culture area always showed statistically greater phytoplankton biomass and nutrient loadings than in adjacent waters, suggesting that this was the “trigger area” of the bloom. The spatial and temporal distribution of diverse HABs in Daya Bay, their ecological characteristics, and their environmental impacts are also discussed in this paper.     

秀丽白虾(Exopalaemon modestus)对亚热带湖库春夏季水体营养水平和浮游生物群落的影响——基于微宇宙模拟实验

秀丽白虾(Exopalaemon modestus)是我国大型淡水湖库的重要经济渔获对象,也是水体中浮游动物的捕食者之一.以往的研究从渔业的角度揭示了秀丽白虾的生物学特征与食性,但少有研究探讨秀丽白虾这一无脊椎动物捕食者对春、夏季浮游生物群落的影响.本研究设计了一个单因素户外微宇宙实验,比较了春季—初夏(4-6月)时期,有无秀丽白虾条件下水体氮、磷浓度以及浮游生物生物量和群落结构的变化.结果 表明:1)秀丽白虾的存在显著降低了春季水体总氮(TN)浓度,增加了水体总磷(TP)浓度,降低了水体TN∶TP比值;2)秀丽白虾的捕食减少了大型枝角类Daphnia pulex和浮游动物总生物量,D.pulex在浮游动物群落中的优势度被轮虫取代,浮游动物群落趋于小型化;3)秀丽白虾显著增加了春季水体叶绿素a浓度,但对浮游植物群落结构的影响不明显.研究表明在大型湖库中,秀丽白虾等无脊椎动物捕食者可能是影响浮游生物群落变化的因素之一,在春季浮游生物研究中应予以关注.     

Lake Kinneret phytoplankton: Stability and variability during twenty years (1970–1989)

The general features of phytoplankton seasonal succession, abundance and distribution in Lake Kinneret, as based on observations from 1970 through 1989, are summarised. Throughout this period of observation, the large, thecate dinoflagellatePeridinium gatunense formed an annual, late winter to early spring bloom resulting in very high standing stock levels. The dominance of these dinoflagellates has a profound impact upon the lake ecosystem. In the summer and fall the phytoplankton assemblage consisted mainly of nanoplanktonic green and blue-green algae and diatoms. Picophytoplankton, mainly picocyanobacteria, were present in low numbers during the dinoflagellate bloom but reached maximum abundance (10⁵ cells · ml^–1) in the epilimnion during the summer and fall. Within a given year, chlorophyll concentrations correlated well with estimates of wet weight biomass, derived from microscope counts. However, interannual averages of chlorophyll did not correlate closely with those for wet weight biomass. Both wet weight biomass and chlorophyll standing stocks fluctuated more than 2 fold from 1970 through 1989 but no extreme, long-term, continuous trend of increase or decrease was observed. Thus, phytoplankton has remained relatively stable although there has been a significant rise in the levels of summer-fall biomass since 1981. The main factor responsible for this may have been increased available phosphorus; the abundance of phytoplankton did not show any clear, long-term relation to that of herbivorous zooplankton.     

Modeling the timing of spring phytoplankton bloom and biological production of the Gulf of St. Lawrence (Canada): Effects of colored dissolved organic matter and temperature

The effects of colored dissolved organic matter (CDOM) from freshwater runoff and seasonal cycle of temperature on the dynamic of phytoplankton and zooplankton biomass and production in the Gulf of St. Lawrence (GSL) are studied using a 3-D coupled physical-plankton ecosystem model. Three simulations are conducted: (1) the reference simulation based on Le Fouest et al. (2005), in which light attenuation by CDOM is not considered and maximum growth rate (_μmax${\mu }_{\max }$) of phytoplankton and zooplankton are not temperature-dependent (REF simulation); (2) light attenuation by CDOM is added to REF simulation (CDOM simulation); and (3) in addition to CDOM, the _μmax${\mu }_{\max }$ of phytoplankton and zooplankton are regulated by temperature (CDOM+TEMP simulation). CDOM simulation shows that CDOM substantially reduces phytoplankton biomass and production in the Lower St. Lawrence Estuary (LSLE), but slightly reduces overall primary production in the GSL. In the LSLE, the spring phytoplankton bloom is delayed from mid-March to mid-April, resulted from light attenuation by CDOM. The CDOM+TEMP simulation shows that the spring phytoplankton bloom in the LSLE is further delayed to July, which is more consistent with observations. Annual primary production is reduced by 33% in CDOM+TEMP simulation from REF and CDOM simulations. Zooplankton production is the same in all three simulations, and export of organic matter to depth is reduced in CDOM+TEMP simulation, suggesting that temperature controlled growth of phytoplankton and zooplankton enhances the coupling between primary production and zooplankton production under the seasonal temperature cycle of the GSL.     

Seasonal and interannual variation of the phytoplankton and copepod dynamics in Liverpool Bay

The seasonal and interannual variability in the phytoplankton community in Liverpool Bay between 2003 and 2009 has been examined using results from high frequency, in situ measurements combined with discrete samples collected at one location in the bay. The spring phytoplankton bloom (up to 29.4 mg chlorophyll m⁻³) is an annual feature at the study site and its timing may vary by up to 50 days between years. The variability in the underwater light climate and turbulent mixing are identified as key factors controlling the timing of phytoplankton blooms. Modelled average annual gross and net production are estimated to be 223 and 56 g C m⁻² year⁻¹, respectively. Light microscope counts showed that the phytoplankton community is dominated by diatoms, with dinoflagellates appearing annually for short periods of time between July and October. The zooplankton community at the study site is dominated by copepods and use of a fine mesh (80 μm) resulted in higher abundances of copepods determined (up to 2.5 × 10⁶ ind. m⁻²) than has previously reported for this location. There is a strong seasonal cycle in copepod biomass and copepods greater than 270 μm contribute less than 10% of the total biomass. Seasonal trends in copepod biomass lag those in the phytoplankton community with a delay of 3 to 4 months between the maximum phytoplankton biomass and the maximum copepod biomass. Grazing by copepods exceeds net primary production at the site and indicates that an additional advective supply of carbon is required to support the copepod community.     

Spatial and Temporal Patterns of Plankton Assemblage Structure in a High Altitude Saline Lake,Namuka Co in Northern Tibet,China

The seasonal variations in biomass, abundance, and species composition of plankton in relation to hydrography were studied in saline lake Namuka Co, northern Tibet, China. The sampling was carried out at approximately monthly intervals from June 2001 to July 2002. The salinity ranged from 5.5 to 26 g/L. The mean annual air and water temperature showed a clear seasonal pattern, which was approximately 4.4 and 7.4°C, respectively, with the lowest water temperature in winter (from December to March, −1°C) and the highest in June and July (18°C). The results showed that 36 taxa of phytoplankton and 16 taxa of zooplankton were identified. Both the biomass and abundance of total phytoplankton were lower in the winter and peaked once or twice during the summer and spring in the early August (8.23 mg/L and 158.2 × 10⁶ ind./L). The seasonal variation in total zooplankton biomass and abundance was characterized by lower values in both winter and early spring, and one maximum (90.5 mg/L and 935 ind. L⁻¹) occurred in the late summer. Major phytoplankton species were Gloeothece linearis, Oscillatoria tenuis, Gloeocapsa punctata, Ctenocladus circinnatus, Ulothrix sp., and Spirogyra sp. And major zooplankton species included Vorticella campanula, Brachionus plicatilis, Daphniopsis tibetana, Cletocamptus dertersi, Arctodiaptomus stewartianus. The production of D. tibetana was 420.3 g m³ a⁻¹. The total number of plankton species has a significant negative correlation with the salinity.     

Feeding activity of adult roach (Rutilus rutilus (L.)), perch (Perca fluviatilis L.) and ruffe (Gymnocephalus cernuus (L.)) in eutrophic Lake Aydat (France)

The feeding activity of adult roach (Rutilus rutilus), perch (Perca fluviatilis) and ruffe (Gymnocephalus cernuus) was studied over a one year cycle to show trophic relations between fish and the prey communities in eutrophic Lake Aydat.Daily consumption rates were referred to the stock of the different age classes of fish. Seven food compartments (phytoplankton, macrophytes, Cladocera, Copepoda, macroinvertebrates, fish and sediment) were used by the fish. The main prey compartment ofR. rutilus were macroinvertebrates (39.7%), sediment (29.0%), macrophytes (15.4%) and phytoplankton (14.9%). Little zooplankton was consumed (2.4%).P. fluviatilis fed mainly on macroinvertebrates and fish fry (both 41.8%) and some zooplankton (15.0%). ForG. cernuus, macroinvertebrates dominated in the diet (97.4%). During spring, Bacillariophyceae and macroinvertebrates represented the principal preys. During summer, insect larvae, phytoplankton and macrophytes were the principal components of the diet. At the end of summer, macrofauna was scare and fish tended to move to the pelagic zone feeding on plankton. In autumn, the prey spectrum was large. During winter, sediment was the most important item consumed, together with some macroinvertebrates. The impact of this fish community through consumption is highest on macroinvertebrates (40.7%), flora (27.7%) and sediment (28.4%). The predation of these three fish on zooplankton was low (1% per day of the zooplankton biomass) and restricted to periods when the other items were scarce. Interspecific competition is not excluded for macroinvertebrates.     

鳑鲏与河蚌交互作用对浮游生物和底栖生物的影响分析

放养河蚌,提高水体透明度以促进沉水植物生长,是湖泊生态修复中的常用手段之一.而小型杂食性鱼类鳑鲏依赖河蚌繁殖,河蚌放养可能会促进鳑鲏种群的发展;而鳑鲏与河蚌交互作用对水生态系统的影响仍研究较少.于2018年11-12月通过原位受控实验,设置对照组、河蚌组、鳑鲏组和河蚌+鳑鲏组,研究了鳑鲏(大鳍鱊Acheilognath...     

Asynchrony of spring phytoplankton response to temperature driver within a spatial heterogeneity bay of Three-Gorges Reservoir, China

The increased air temperature is expected to have important driver on spring phytoplankton dynamics. To test whether spatial heterogeneity modifies the synchronous responses of phytoplankton to regional temperature driver, we evaluate temporal coherences for physical factors (temperature, water stability and non-algal light extinction), nutrients (nitrogen, phosphorus and silicon), and biomass and density of phytoplankton by Pearson correlation analysis and synchrony for phytoplankton community dynamics by Mantel test and nonmetric multi-dimensional scaling (NMS), during spring bloom (February 23–April 28, 2005) within Xiangxi Bay, a high spatial gradient bay of Three-Gorges Reservoir (China). The high level of temporal coherences for surface water temperature (r = 0.946, p < 0.01) and relative water column stability (r = 0.750, p < 0.01) were found between pair sites (A and B), in which the increase trends occurred with increase in regional air temperature during the study period. However, the low synchrony for phytoplankton dynamics were indeed observed between Site A and B, especially for the density of common dominant taxa (Cyclotella spp.: r = 0.155, p = 0.388) and community structure (Mantel test: r = 0.351). Moreover, the local habitat characteristics such as nutrient (nitrogen and phosphorus) and non-algal light extinction showed low levels of temporal coherence. It indicated that local community of phytoplankton varies rather independently within the single lentic bay with high spatial heterogeneity and that dispersal of algal organisms among locations cannot overwhelm out these local dynamics. Contrary to many studies, the present results argued that, in a small geographic area (i.e., a single reservoir bay of approximately 24 km length), spatial gradients also may influence spring phytoplankton response to regional temperature driver.     

Size-fractionated phytoplankton biomass,primary production and respiration in the Nervión–Ibaizabal estuary: A comparison with other nearshore coastal and estuarine ecosystems from the Bay of Biscay

The seasonal pattern of size-fractionated phytoplankton biomass, primary production and respiration was investigated along the longitudinal axis of the Nervión–Ibaizabal estuary (Bay of Biscay) from April 2003 to September 2004. Environmental factors influencing phytoplankton dynamics were also studied. Chlorophyll a biomass showed a longitudinal pattern of increase from the outer Abra bay to the inner estuary. On a seasonal scale, in the intermediate and inner estuary phytoplankton biomass maxima were registered in summer, the warmest and driest season, whereas in the outer bay chlorophyll a peaks occurred in May 2004, but were delayed to August 2003, likely due to a very rainy spring. Data suggest that river flow exerts a marked influence on the timing of phytoplankton biomass maxima in this estuary, decreased river flows providing a lowering of turbidity and an increase in water residence time needed for chlorophyll a to build up. Nutrient concentrations were high enough not to limit phytoplankton growth throughout the annual cycle, except silicate and occasionally phosphate in the outer bay during summer. Silicate concentration correlated positively with river flow, whereas ammonium and phosphate maximum values were generally measured in the mid-estuary, suggesting the importance of allochthonous anthropogenic sources. In the intermediate and inner estuary phytoplankton biomass was generally dominated by >8 μm size-fraction (ca. 60%), but in August 2003 <8 μm size-fraction increased its contribution in the intermediate estuary. It is argued that the lower nutrient concentrations measured in August 2003 than in August 2004 could have played a role. This is the first study in which phytoplankton primary production rates have been measured along the longitudinal axis of the Nervión–Ibaizabal estuary. Throughout the annual cycle these rates ranged from 0.001 to 3.163 g C m^?3 d^?1 and were comparable to those measured in nearby small estuaries of the Basque coast and other larger estuaries on the Bay of Biscay. Surface plankton community respiration rate maxima were measured during the spring 2004 chlorophyll a peak in the Abra bay and in summer months at the mid and inner estuary, coinciding with chlorophyll a biomass and primary production maxima. In general, respiration rates showed a positive correlation with temperature. In order to compare results from the Nervión–Ibaizabal estuary with other nearshore coastal and estuarine ecosystems within the Bay of Biscay a review of existing information on phytoplankton biomass and primary production dynamics was performed.     

江汉平原长湖浮游植物初级生产力的季节性变化及其驱动因子

为探究长江中下游富营养化浅水湖泊的浮游植物初级生产力季节性演替特征及其驱动因子,本研究于2020年4月(春)、8月(夏)、10月(秋)及2021年1月(冬)对湖北长湖浮游植物进行采样调查,同时运用黑白瓶测氧法及VGPM模型估算法分别估算了其浮游植物生产力水平,并探究驱动初级生产力季节性变化的主要环境因子。结果显示,4个季节共鉴定出浮游植物194种,其中绿藻门(95种,49%)和硅藻门(40种,21%)居绝对优势地位;黑白瓶法测得浮游植物水柱总生产力(P_t)季节变化为:夏季((1841.24±345.93) mg C/(m²·d))>秋季((1324.14±208.34) mg C/(m²·d))>春季((847.50±247.72) mg C/(m²·d))>冬季((711.43±133.52) mg C/(m²·d)),其中M2站位在夏季采样时(2424.66 mg C/(m²·d))水柱总生产力最高;在垂直空间上,浮游植物总生产力(G...     

An evaluation of the role of daphnids in controlling phytoplankton biomass in clear water versus turbid shallow lakes

Phytoplankton and zooplankton were monitored during 2 years in four eutrophic shallow lakes (two turbid and two clear water) from two wetland reserves in Belgium. In each wetland, phytoplankton biomass was significantly higher in the turbid lake than in the clear water lake. Although total macrozooplankton biomass and the contribution of daphnids to total zooplankton biomass was comparable in the clear water and the turbid lakes, the grazing pressure of macrozooplankton on phytoplankton as estimated from zooplankton to phytoplankton biomass ratios was higher in the clear water lakes. Estimated grazing by daphnids in the clear water lakes was always high in spring. In summer, however, daphnid biomass was low or daphnids were even absent during prolonged periods. During those periods phytoplankton was probably controlled by smaller macrozooplankton or by submerged macrophytes through nutrient competition, allelopathic effects or increased sedimentation rates in the macrophyte vegetation.     

Influences of the Gulf of Maine intrusion on the Massachusetts Bay spring bloom: A comparison between 1998 and 2000

The Boston Harbor, Massachusetts Bay and Cape Cod Bay system (MBS) is a semi-enclosed coastal embayment located in the western Gulf of Maine (GOM). The strength of the spring bloom in the MBS varies dramatically and the underlying mechanisms are not well understood. It has been hypothesized that the weak (or missing) 1998 spring bloom was due to increased zooplankton grazing pressure after a relatively warmer winter that led to earlier development of zooplankton populations. However, chlorophyll concentrations were low in the entire GOM region during the spring bloom period of 1998. These low chlorophyll waters would enter the MBS by persistent intrusion from the GOM and affect the MBS spring bloom by reducing the accumulation of phytoplankton biomass, suggesting an alternative explanation for the missing bloom. In this study, the influences of the intruding GOM waters on the MBS spring bloom are examined using numerical simulations and observations to compare two contrasting spring blooms (a weak bloom in 1998 versus a strong bloom in 2000). The results indicate that intruding waters from the GOM significantly reduced the strength of spring bloom in 1998, but enhanced the spring bloom in 2000. A theoretical analysis suggests that the influence of the intrusion on the MBS spring bloom is mostly limited to the northern portion of the MBS, and the impact area is determined by local net growth, velocity of intruding flow and horizontal mixing. The intrusion of GOM waters carries the signal of long-term variability in the GOM region and thus may be an important oceanic pathway for climate changes to impact the MBS ecosystem.     

Phosphorus and nitrogen limitation of algal biomass across trophic gradients

Regression results based on data from 46 northern temperate lakes show that total phosphorus (TP) is the best predictor for phytoplankton (as chl-a) at lower trophic levels, TP < 200 mg · m^–3. A regression including both TP and TN as regressors is the best predictor for lakes with TP > 200 mg · m^–3. However, the good correlation is probably due to a high correlation between lake average chl-a (all years observed) and lake average TP and TN. Within single hypereutrophic lakes, TN alone is the best predictor. It was not possible to identify a medium trophic domain where TN and TP in combination was the best predictor for chl-a. The ratio TN:TP in the water decreases from about 40 to about 5 with increasing trophic level. Optimum TN:TP ratio for algal species with high abundance during late summer and autumn reflects this decreasing ratio, but within a lesser range, i.e., 20 to 5. In contrast, TN:TP ratios for species abundant during the early vernal period showed no, or an inverse, relation to the TN:TP ratio of the water.     

Differences in zooplankton feeding rates and isotopic signatures from three temperate lakes

Bacteria and phytoplankton are integral in the mobilization and transfer of organic matter to higher trophic levels. Hence, we examined their role in zooplankton diets and assessed trends in their nitrogen isotopic variability. We performed feeding experiments with natural particulate organic matter (POM) and four zooplankton groups (Daphnia, Holopedium, large calanoids and small calanoids) to (1) examine whether there are differences in consumption (presented as clearance and ingestion rates) of phytoplankton and bacteria, and (2) determine whether differences in zooplankton clearance and ingestion rates are correlated with their δ¹⁵N isotopic signatures. In general, phytoplankton and bacteria clearance rates and biomass ingested per animal varied significantly among different zooplankton groups within lakes and between lakes for a given zooplankton group. Within a given lake, Daphnia and Holopedium had the highest phytoplankton and bacteria clearance and ingestion rates, followed by large calanoids, and then small calanoids. For a given zooplankton group, bacteria and phytoplankton clearance rates varied among lakes. In contrast, phytoplankton ingestion rates were consistently highest in Dickie Lake for all taxa, whereas bacteria ingestion rates were more variable among lakes for the different zooplankton taxa. The percentage contribution of different phytoplankton taxa to the biomass of phytoplankton ingested also varied significantly among lakes for a given taxa, but there were few differences within a given lake among zooplankton. Zooplankton δ¹⁵N_DOMC values were correlated with their size adjusted phytoplankton and bacteria clearance and ingestion rates. The correlations were stronger with (1) phytoplankton compared to bacteria, and (2) clearance rates compared to ingestion rates of biomass. Together our results suggest that zooplankton taxa with low phytoplankton and bacteria clearance and ingestion rates and higher δ¹⁵N_DOMC are likely exploiting food sources from higher trophic levels.