Salinity, light, and chlorophyll-a in the Hunter River Estuary Brian G. Sanderson and Anna M. Redden January 14, 2006 Contents 1 Introduction 2 2 Field Excursions 3 3 Results 4 3.1 24/7/05 Fluorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3.2 9/8/05 Fluorescence, phytoplankton counts, nutrients, chlorophyll-a . . . . . . . . 10 3.3 12/8/05 Dilution experiment: grazing and phytoplankton growth . . . . . . . . . . 22 3.4 16/8/05 Zooplankton trawls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 4 Discussion 28 1 Light and chlorophyll-a, Sanderson 1 2 Introduction The NSW Integrated Monitoring of Environmental Flows program requires information about planktonic processes in the Hunter River Estuary. Specific planktonic processes that have been measured and reported here include: • Results from dilution experiments that determine potential phytoplankton growth rates and zooplankton grazing rates. • Dilution experiments were also used to determine the extent that nutrients limit phytoplankton growth. The present work also reports additional measurements from 4 field trips into the Hunter River Estuary that supported the above experimental measures of planktonic processes. In particular, the present work will document: • Salinity, temperature, and turbidity observations made along the length of the Hunter River Estuary on each of the 4 sampling days. • Estimates for chlorophyll-a were obtained from fluorescence measurements made along the length of the estuary. • Phytoplankton counts and zooplankton counts are reported at locations along the length of the estuary. • Light profiles were measured at locations along the length of the estuary. • Nutrient concentrations were measured along the length of the estuary. Relationships between salinity and chlorophyll-a will be examined. Light measurements are used to determine the light attenuation coefficient κ which is important for understanding light limitation of primary production in the estuary. Relationships between turbidity and κ is established. This may be useful for future work since turbidity measurements are commonly available. Light and chlorophyll-a, Sanderson 2 3 Field Excursions Four field excursions were undertaken: • On 24/7/05 near-surface in-situ fluorescence (from which chlorophyll-a is estimated) was measured along the length of the estuary along with profiles of salinity, temperature, turbidity, and dissolved oxygen. The dissolved oxygen sensor was probably reading low, but spatial structure in the measurements is still represented well. The primary purpose of these measurements was to provide background information for designing the following program of biological sampling and experiments. • More extensive sampling was undertaken on 9/8/05. Near-surface in-situ fluorescence measurements were made in conjunction with salinity, temperature, turbidity, and dissolvedoxygen profiling. Light intensity was measured as a function of depth at 9 stations scattered along the length of the estuary. Samples were taken for nutrient analysis, chlorophyll extractions, and for obtaining phytoplankton counts. • On 12/8/05 water samples were obtained from 3 locations in order to undertake dilution experiments to measure: grazing by zooplankton, whether or not nitrogen was limiting, and potential phytoplankton growth rates when light is not limiting. Here we also report the salinity, temperature, and turbidity observations made concurrent with the collection of water samples for the dilution experiment. • On 16/8/05 zooplankton tows were made at 4 stations scattered along the length of the estuary. These measurements were taken after dark. Here we also report concurrent measurements of temperature, salinity and turbidity in order to provide contextual information. Light and chlorophyll-a, Sanderson 3 4 Results 3.1 24/7/05 Fluorescence Salinity, temperature, dissolved oxygen, and turbidity profiles were measured along the length of the Hunter River Estuary (Figure 1). Temperature was about 16 o C near the mouth of the estuary and fell to 12 o C 40 km upstream at the limit of salt intrusion. The seasonal cycle of heating and cooling results in cooler conditions inland during the winter. The along-channel temperature gradient is reversed in summer. Baroclinic circulation (leading to vertical gradients) is driven mostly by the horizontal density gradient associated with salinity. In winter the temperature gradient partially reduces the density gradient due to salinity. In summer temperature effects whereas is reinforces the density gradient due to salinity. Given the surface-layer of the ocean is usually well-mixed to depths associated with the estuary, it follows that vertical gradients of salinity are small in the lower estuary. Similarly, there is no salt in the upper estuary so vertical salinity gradients must also be small there. In between, baroclinicity associated with the horizontal density gradient causes more dense (saline) water to slump beneath less dense water. This mechanism constantly generates vertical salinity gradients which are, in turn, eroded by vertical mixing associated with wind and tide. Salinity typically varies by about 3 ppt through the water column. In the upper estuary, there is clear vertical stratification of temperature — although this is typically ∼0.25 o C from top to bottom. Figure 2 shows the vertically-averaged salinity, temperature, turbidity, and dissolved oxygen along the length of the estuary. Sampling was undertaken a long way up the estuary to positions where the water was essentially fresh. Temperature and salinity decrease upstream whereas turbidity and dissolved oxygen increased upstream. Figure 3 shows chlorophyll-a increases upstream consistent with increased dissolved oxygen that may be associated with primary production. Plotting chlorophyll-a against salinity (Figure 4), it is clear that the high upstream chlorophyll-a near the head of the estuary is decoupled from downstream areas that have higher salinity. If chlorophyll-a was conserved as it mixed downstream then there would be a more linear rela- Light and chlorophyll-a, Sanderson 5 tionship between salinity and chlorophyll-a (assuming steady-state). The present measurements indicate upstream phytoplankton suffer severe losses as they are mixed downstream into more salty water. Figure 5 shows salinity profiles near the junction of a creek draining Korangang. The main channel is vertically well-mixed upstream of the junction. Water draining from Korangang is relatively fresh. This illustrates the way in which horizontal mixing can result from oscillatory tidal currents and channel junctions. This particular side creek is not explicitly treated in the hydrodynamic model used for salinity calculations — rather it is parameterized as a horizontal eddy-diffusivity. Light and chlorophyll-a, Sanderson 6 Salinity Hunter River estuary, 24/07/2005 Depth (m) 0 −5 20 30 SB −10 0 5 10 HB 10 RT Hunter River 15 20 25 Distance up estuary (km) 30 35 40 Temperature Hunter River estuary, 24/07/2005 Depth (m) 0 14 −5 12 16 SB −10 0 5 HB 10 RT Hunter River 15 20 25 Distance up estuary (km) 30 35 40 DO, percent saturated Hunter River estuary, 24/07/2005 Depth (m) 0 −5 70 75 SB −10 0 5 80 HB 10 15 20 25 Distance up estuary (km) RT Hunter River 30 35 40 Turbidity Hunter River estuary, 24/07/2005 Depth (m) 0 −5 −10 0 5 10 20 3050 SB 5 HB 10 15 20 25 Distance up estuary (km) RT Hunter River 30 35 40 Figure 1: Spatial structure of salinity, temperature, dissolved oxygen, and turbidity on 24/7/05. Light and chlorophyll-a, Sanderson 7 TURB (NTU) 60 40 20 0 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 45 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 45 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 45 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 45 40 S (ppt) 30 20 10 0 T (Celcius) 17 16 15 14 13 12 DO (%) 85 80 75 70 Figure 2: Along-channel distribution of vertically-averaged: turbidity, salinity, temperature, and dissolved oxygen on 24/7/05. Light and chlorophyll-a, Sanderson 8 Chl−a (µg/l) 15 10 5 0 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 15 TURB (NTU) 10 5 0 −5 Figure 3: Along-channel distribution of near-surface chlorophyll-a on 24/7/05. 15 Chlorophyll−a (µg/l) Chlorophyll−a (µg/l) 15 10 5 0 0 10 20 Salinity (ppt) 30 10 5 0 0 10 20 30 Turbidity (NTU) 40 50 Figure 4: Relationships between chlorophyll-a and salinity, and chlorophyll-a and turbidity on 24/7/05. Light and chlorophyll-a, Sanderson 9 A 0 −1 −2 B −3 A −4 C −5 15 20 25 T ( C), S (ppt) 30 o 0 0 −1 −1 −2 −2 −3 −3 C −4 −5 15 20 25 o T ( C), S (ppt) B −4 30 −5 15 20 25 T ( C), S (ppt) 30 o Figure 5: Salinity profiles where a creek draining Korangang runs into the Hunter River Estuary after about an hour of outgoing tide. Profile A is taken in the Hunter River Estuary upstream of the junction. Profile B is taken in the Hunter River Estuary downstream of the junction. Profile C is taken in the side creek. Light and chlorophyll-a, Sanderson 3.2 10 9/8/05 Fluorescence, phytoplankton counts, nutrients, chlorophylla Vertical gradients are weaker in Figure 6. These measurements of salinity, temperature, and turbidity are similar to those obtained on the previous survey. Measurements were not made so far upstream on this occasion, however, so the lowest salinities in Figure 7 are still markedly above those of freshwater. Thus, although Figure 8 shows chlorophyll-a higher upstream, it does not get as high as measurements made 24/7/05. Phytoplankton counts were made using two replicates at each of the 9 measurement sites. The total number of phytoplankton is plotted as a function of chainage in Figure 8. The large increase in chlorophyll-a near the head of the estuary is reflected by large phytoplankton counts, but otherwise chlorophyll-a and total phytoplankton count are poorly related (Figure 9). Given that phytoplankton can have vastly different sizes, it is hardly surprising that total phytoplankton counts are a poor estimate of the amount of bulk measures of a phytoplankton community (like chlorophyll-a). Table 1 documents the distribution of phytoplankton counts among various taxonomic groups. The very high diatom counts at the upstream site are due to Aulacoseira. Figure 10 shows that Aulacoseira sp. counts vary exponentially over chainages 20-35 km upstream from the mouth of the estuary. Aulacoseira is a common freshwater species in the Hunter River and it is not surprising that its concentration drops by a factor of 3 for every 3.7 km of displacement into more saline water. It would seem that Aulacoseira suffer mortality when salinity increased and the distribution of Aulacoseira could be well represented using a mixing model. The number of species (Table 1) is in the range 15-18 throughout except at the most upstream sites. High counts of Aulacoseira sp. at the upstream sites are expected to bias the number of species to lower values, as observed. Overall, it seems that the species richness does not vary greatly along the length of the estuary for which measurements were made. Different groupings of phytoplankton vary substantially along the length of the estuary (Table 1 and Figure 11). Diatoms (Bacillariophyceae) are most abundant near the head of the estuary where there are large numbers of freshwater Aulacoseira. Counts of diatoms are low in mid-estuary Light and chlorophyll-a, Sanderson 11 Salinity Hunter River estuary, 09/08/2005 Depth (m) 0 30 −5 −10 0 SB 5 10 20 HB 10 RT Hunter River 15 20 25 Distance up estuary (km) 30 35 40 Temperature Hunter River estuary, 09/08/2005 Depth (m) 0 16 −5 −10 0 SB 5 HB 10 RT Hunter River 15 20 25 Distance up estuary (km) 30 35 40 DO, percent saturated Hunter River estuary, 09/08/2005 Depth (m) 0 −5 −10 0 75 SB 5 HB 10 RT Hunter River 15 20 25 Distance up estuary (km) 30 35 40 Turbidity Hunter River estuary, 09/08/2005 Depth (m) 0 5 −5 −10 0 10 SB 5 HB 10 15 20 25 Distance up estuary (km) RT Hunter River 30 35 40 Figure 6: Spatial structure of salinity, temperature, dissolved oxygen, and turbidity on 9/8/05. Light and chlorophyll-a, Sanderson 12 TURB (NTU) 20 15 10 5 0 0 5 10 15 20 Distance up estuary (km) 25 30 35 0 5 10 15 20 Distance up estuary (km) 25 30 35 0 5 10 15 20 Distance up estuary (km) 25 30 35 0 5 10 15 20 Distance up estuary (km) 25 30 35 40 S (ppt) 30 20 10 0 17 T (ppt) 16.5 16 15.5 15 14.5 DO (%) 80 78 76 74 Figure 7: Along-channel distribution of vertically-averaged: turbidity, salinity, temperature, and dissolved oxygen on 9/8/05. Light and chlorophyll-a, Sanderson 13 Total phytoplankton (#/ml) 10000 8000 6000 4000 2000 0 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 10 Chl−a (µg/l) 8 6 4 2 0 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 0 5 10 15 20 25 Distance up estuary (km) 30 35 40 10 TURB (NTU) 8 6 4 2 0 Figure 8: Top, along-channel distribution of total phytoplankton count on 9/8/05. Middle, alongchannel distribution of near-surface chlorophyll-a on 9/8/05. Bottom, turbidity. Light and chlorophyll-a, Sanderson 14 9000 8000 Total phytoplankton (#/ml) 7000 6000 5000 4000 3000 2000 1000 0 0 2 4 6 Chlorophyll−a (µg/l) 8 10 Figure 9: Total phytoplankton count is poorly related to chlorophyll-a on 9/8/05. Light and chlorophyll-a, Sanderson chainage salinity 15 Bacillario- Dino- Chrysophyceae Chloro- phyceae phyceae +Chryptophyceae phyceae Un-id Total no. nano species +Euglenophyceae +Prasinophyceae +Prasinophyceae km ppt #/ml #/ml #/ml #/ml #/ml #/ml 34.3 2.8 7047 (6605) 0 236 885 177 8345 10 32.0 4.0 3389 (3140) 0 74 575 162 4199 10 28.4 6.9 915 (778) 0 171 142 42 1269 14 26.1 10.2 664 (551) 127 365 101 165 1420 15 22.8 14.7 441 (264) 195 420 58 528 1641 16 20.6 19.0 243 (96) 147 611 8 774 1782 17 17.1 22.9 433 (0) 217 541 10 1022 2222 18 11.5 30.3 1396 (0) 176 639 0 776 2986 17 4.5 32.8 1781 (0) 50 393 0 246 2469 18 Table 1: Phytoplankton counts of various groups. The bracketed numbers under Bacillariophyceae are counts of Aulacoseira sp. which are responsible for the high counts at the upstream sites. Cyanophyceae counts were zero at all sites. but increase significantly near the mouth due to the presence of Thalassiosira (Figure 11a). Thus, diatoms seem to be represented by both oceanic and freshwater species. Green algae are abundant upstream but counts fall as the salinity increases (Figure 11b) so one might conclude that the green algae are also associated with freshwater. Dinoflagellates and unidentified nanoplankton are low near the mouth and head of the estuary but high in mid-estuary. This indicates that the Dinoflagellates and the unidentified nanoplankton are estuarine in origin — so they grow within the estuary whereas the green algae and diatoms are mostly associated with boundary conditions. Figures 12 and 13 show a large number of light profiles made at various distances (chainage) upstream from the mouth of the estuary. Exponential functions are fitted to each profile to Light and chlorophyll-a, Sanderson 16 4 10 Aulacoseira (#/ml) 3 10 2 e−folding scale = 3.4 km 10 1 10 20 25 30 35 chainage (km) Figure 10: Aulacoseira sp. decay with distance downstream. The concentration changes by a factor of 3 every 3.7 km along-channel. Light and chlorophyll-a, Sanderson 17 8000 1000 (a) green algae diatoms 4000 2000 0 (b) 800 6000 600 400 200 0 10 20 30 0 40 0 10 Chainage (km) (c) 200 150 100 50 0 0 10 20 30 40 40 1000 800 600 400 200 0 Chainage (km) Figure 11: 30 (d) 1200 unID nonoplankton dinoflagellates 250 20 Chainage (km) 0 10 20 30 40 Chainage (km) Along-channel distribution of phytoplankton groups. In plot (a) Aulacoseira sp. make up most of the counts near the head of the estuary whereas Thalassiosira sp. make up most of the counts at the mouth. The red crosses in plot (d) are a composite of Chrysophyceae, Chryptophyceae, Euglenophyceae, and Prasinophyceae. Light and chlorophyll-a, Sanderson 18 determine surface light intensity and the light attenuation coefficient κ. The e-folding scale for light is given by κ−1 which is the depth (m) of water required to attenuate the light intensity by a factor of 1/e ∼ 0.37. Light attenuation κ increases progressing upstream. Upstream, the e-folding scale is 0.5 m. This means that the surface light intensity reduces by a factor of 0.0025 at a depth of 3 m. High chlorophyll-a concentrations in the upstream waters would seem to require some buoyancy mechanism to stabilize the water column — perhaps the temperature gradient. Figure 14 plots light attenuation against chlorophyll-a and also against turbidity. While chlorophyll-a contributes to light attenuation, it is not the dominant factor. There is a clear relationship between light attenuation and turbidity. Light and chlorophyll-a, Sanderson 19 Chainage = 4.5 km Chainage = 11.506 km 0 0 1200 hrs 1315 hrs −0.5 z (m) z (m) −0.5 −1 −1.5 −1 −1.5 I = 1246EXP(0.603z) −2 0 500 1000 2 Light (W/m ) I = 1624EXP(1.19z) −2 1500 0 500 1000 2 Light (W/m ) Chainage = 17.128 km Chainage = 20.58 km 0 0 1400 hrs 1430 hrs −0.5 z (m) z (m) −0.5 −1 −1.5 −1 −1.5 I = 877EXP(1.01z) −2 0 200 400 Light (W/m2) I = 1366EXP(1.15z) −2 600 800 0 Chainage = 22.763 km 500 1000 Light (W/m2) 1500 Chainage = 25.399 km 0 0 1500 hrs 1535 hrs −0.5 z (m) −0.5 z (m) 1500 −1 −1.5 −1 −1.5 I = 1078EXP(1.32z) −2 0 200 400 600 2 Light (W/m ) 800 I = 996EXP(1.41z) −2 1000 0 200 400 2 Light (W/m ) 600 800 Figure 12: Light profiles, along with fitted curves based on a best-fit light attenuation coefficient and surface irradiance. The coefficient of light attenuation increases with distance from the estuary mouth (chainage). Light and chlorophyll-a, Sanderson 20 Chainage = 27.07 km Chainage = 31.919 km 0 0 1605 hrs 1635 hrs −0.5 z (m) z (m) −0.5 −1 −1.5 −1 −1.5 I = 783EXP(1.53z) −2 0 200 400 2 Light (W/m ) 600 I = 335EXP(1.92z) −2 800 0 100 200 2 Light (W/m ) 300 Chainage = 34.174 km 0 1705 hrs z (m) −0.5 −1 −1.5 I = 279EXP(1.96z) −2 0 100 200 2 Light (W/m ) 300 Figure 13: Light profiles, along with fitted curves based on a best-fit light attenuation coefficient and surface irradiance. 10 8 8 Chlorophyll−a (µg/l) 10 6 4 2 0 0 10 20 Salinity (ppt) 6 4 2 0 30 2 2 1.5 1.5 −1 κ (m ) 21 κ (m−1) Chlorophyll−a (µg/l) Light and chlorophyll-a, Sanderson 1 0.5 0 5 10 15 Turbidity (NTU) 20 1 0.5 κ = 0.242 + 0.116Turb 0 0 2 4 6 Chlorophyll−a (µg/l) 8 10 0 0 5 10 Turbidity (NTU) 15 Figure 14: Relationships between chlorophyll-a and salinity, and chlorophyll-a and turbidity on 9/8/05. Relationships with light attenuation are also shown Light and chlorophyll-a, Sanderson 3.3 22 12/8/05 Dilution experiment: grazing and phytoplankton growth Table 2 presents vertically-averaged properties of the water column at 3 sites where water was collected for dilution experiments. The dilution site with the lowest salinity still had salinity larger than 6. From Figures 4 and 14 it is clear that high chlorophyll-a concentrations are restricted to less saline waters. Chlorophyll-a does not vary greatly in the water samples collected for the dilution experiment. Note, the highest salinity used for dilution experiments was 18.62 ppt. Small filamentous macroalgae were found in more saline water and these can disrupt dilution experiments. If a water sample is incubated under sufficient photosynthetically active radiation then phytoplankton can be expected to grow. Measuring chlorophyll-a before and after the incubation gives an estimate of phytoplankton growth minus any losses due to grazing by zooplankton (apparent phytoplankton growth). If a sample is diluted with filtered seawater, then the grazing will be reduced. Incubating several samples with a range of dilutions gives apparent phytoplankton growth rate as a function of dilution. Theoretically, the apparent phytoplankton growth rate should reduce as the fraction of unfiltered seawater increases. Fitting a linear line to a plot of apparent growth rate against fraction of unfiltered seawater gives a fit where the negative of the slope is an estimate of zooplankton grazing rate and the y-intercept is an estimate of phytoplankton growth rate. Figure 15 shows results for samples from 3 sites at locations documented in Table table:dilution. (Note, location is most logically referenced to salinity in a tidal estuary.) Apparent growth rate ln(chl-t/chl-i)/∆t is plotted against fraction unfiltered seawater. Here, chl-t is the chlorophyll-a concentration after a 24 hour incubation and chl-i is the initial chlorophyll-a concentration. (The incubation period ∆t, being 1 day, is implicit in the y-axis labelling of Figure 15.) Grazing rates g, growth rates µ, and initial chlorophyll-a chl-i are recorded in Table 2. Initial chlorophyll-a was in the range 1.5-3 µg/L, as expected given the salinity of the sampling locations. Growth rates are much higher than grazing rates. Indeed, the average growth rate is 1.23 day −1 which amounts to an increase by a factor of 3.42 in a 1 day period. Subtracting out the effect of grazing (with no dilution) gives an average growth rate of 1.08 day−1 which still amounts to Light and chlorophyll-a, Sanderson 23 chainage salinity temperature DO turbidity comment µ g chl-i (km) (ppt) (Celsius) (%) (NTU) day−1 day−1 µg/L 4.5 32.4 15.4 70 7.1 9.9 28.9 14.0 73 7.6 13.2 23.1 13.7 71 7.1 17.1 16.1 13.5 71 4.0 18.6 18.6 13.8 72 4.1 Dilution 3 1.2 0.17 2.3 22.8 13.6 13.6 70 4.3 24.0 12.6 13.7 71 4.6 Dilution 2 1.2 0.09 1.5 28.4 6.9 13.4 69 7.0 Dilution 1 1.3 0.20 3.0 Table 2: Growth rates µ and grazing rates g from dilution experiments along with contextual physical information. Salinity, temperature, DO, and turbidity are vertically-averaged through the water column. The DO sensor was not calibrated so DO is biased low. DO is essentially the same at all stations. phytoplankton growing by a factor of 2.95 in a 1 day period. On the other hand, fluorescence measurements in the Hunter River Estuary are relatively stable in the period 24/7/05 through 9/8/05 and consistent with initial chlorophyll-a values on 12/8/05. Clearly the growth rates measured by the dilution experiment are potential growth rates whereas something other than grazing is limiting phytoplankton growth in the estuary. Some of the samples had bioavailable nitrogen added before incubation. These samples are plotted in purple in Figure 15. Clearly, samples with added nutrient grew the same as samples for which nutrient was not added. It follows, therefore, that nitrogen is not limiting phytoplankton growth rate. Nitrogen is considered the nutrient most likely to limit growth in Australian estuaries (Harris 199X) so it would seem that nutrients are not limiting phytoplankton growth in the Hunter River Estuary. Growth rates (Table 2) are essentially the same at each of the three locations from which samples were obtained for dilution studies. On the other hand, Table 3 shows that the more Light and chlorophyll-a, Sanderson chainage salinity Bacillariophyceae 24 Dino- Chrysophyceae Chloro- Un-id Total phyceae +Chryptophyceae phyceae nano no. species +Euglenophyceae +Prasinophyceae km ppt +Prasinophyceae 18.6 18.6 2712 (0,2683) 118 265 177 176 3448 10 24.0 12.6 803 (731,24) 24 306 47 165 1345 13 28.4 6.9 1371 (560,206) 133 590 30 542 2666 20 Table 3: Phytoplankton composition in dilution samples. Counts of Cyanophyceae were zero for all dilution samples. The bracketed numbers under Bacillariophyceae represent counts of (Aulacoseira sp., Chaetoceros spp.) saline site was dominated by Chaetoceros spp. which was not abundant at the other sites. It should also be noted that whereas Chaetoceros spp. dominated samples with salinity 18.6 ppt on 12/8/05, they were not abundant in samples with similar salinity on 9/8/05. Clearly there is much spatio-temporal variability in phytoplankton counts that would make them difficult to model. Bulk properties, such as chlorophyll-a, are more readily modelled. 3.4 16/8/05 Zooplankton trawls Table 4 shows vertically-averaged water column properties at sites where zooplankton trawls were done. The most upstream zooplankton trawl site was in water with salinity 3.95 ppt, where chlorophyll-a concentrations can be expected to be somewhat elevated. Filamentous macroalgae were present at the more saline sites, but were not so abundant as on the previous field trips (macro-algae abundance was qualitatively assessed from samples obtained in a small zooplankton net). Table 5 shows zooplankton counts/m3 from four sites along the Hunter River estuary. An ensemble of three tows were made at each site using a 100 µm mesh net. The duration of each tow was 2 minutes. The most marked features in this data are: Light and chlorophyll-a, Sanderson Figure 15: Plots of results from the dilution experiments. 25 Light and chlorophyll-a, Sanderson 26 chainage salinity temperature DO turbidity comment (km) (ppt) (Celsius) (%) (NTU) 9.936 30.71 15.38 74 7.8000 13.190 29.67 15.45 80 9.1667 17.127 24.12 14.89 78 5.0000 18.644 20.23 14.41 78 4.5000 21.469 20.23 14.63 81 5.5857 27.089 14.30 14.37 77 4.5750 28.399 12.27 14.16 73 6.2000 34.289 6.42 14.32 74 8.7000 44.000 3.95 14.53 80 18.3000 Zoop stn 4 Zoop stn 3 Zoop stn 2 Zoop stn 1 Table 4: Contextual information for zooplankton counts. Quantities are vertically-averaged through the water column. • Fish eggs are more abundant in the low salinity waters at the upstream site (site 1). • Calanoid adults were markedly more abundant in the low salinity waters at the upstream site (site 1). • Other copepods (Copepod Nauplii, juvenile Calanoid Copepodites, Cyclopoids, Harpacticoids) were more broadly distributed through the estuary with a tendency for concentration to increase somewhat downstream. • The most downstream site (salinity 30.7 ppt) had markedly high concentrations of Noctiluca, a marine heterotrophic dinoflagellate. Filamentous algae were also abundant at this site. Light and chlorophyll-a, Sanderson Taxa 27 Zoop stn 1 Zoop stn 2 Zoop stn 3 Zoop stn 4 (#/m3 ) (#/m3 ) (#/m3 ) (#/m3 ) Copepod Nauplii 36 ± 4 34 ± 8 34 ± 4 71 ± 30 Calanoid Copepodites (Juveniles) 62 ± 7 15 ± 2 29 ± 2 103 ± 28 Calanoid Adults 243 ± 50 14 ± 1 14 ± 3 8±1 Cyclopoids 0 5±2 16 ± 3 7±3 Harpacticoids 10 ± 0.9 0.67 ± 0.3 18 ± 2 38 ± 8 Noctiluca 0.3 ± 0.3 0.3 ± 0.3 22 ± 6 546 ± 170 Oikopleura (Appendicularian) 3±1 0.3 ± 0.3 1 ± 0.6 0.3 ± 0.3 Polychaete Trochophore/ Larvae 1 ± 0.6 1.3 ± 0.9 36 ± 4 70 ± 30 Bivalve Larvae 0.3 ± 0.3 2±1 2±1 1±1 Gastropod Larvae 4±2 5±3 3.7 ± 2 2.7 ± 1 Fish Eggs 10 ± 2 0.3 ± 0.3 1 ± 0.6 1.3 ± 0.7 Shrimp Larvae 0 0 0 2 ± 0.6 Jellyfish 0 0 0.3 ± 0.3 0 Chironomid 0.3 ± 0.3 0 0 0 Cladoceran 1.3 ± 0.9 0 0.3 ± 0.3 0.67 ± 0.3 Saltwater Mite 0.3 ± 0.3 0 0.3 ± 0.3 0 Crab Zoea 1.3 ± 0.9 0.3 ± 0.3 1 ± 0.6 8±5 Ostracod 0.3 ± 0.3 0 0.3 ± 0.3 0 Barnacle Cypris Larvae 0 0 7.7 ± 3 4±1 Barnacle Nauplii 0 0.67 ± 0.3 181 ± 6 31 ± 15 Snail Egg Case 0 0 0.3 ± 0.3 0 Cumacean 0 0 0.3 ± 0.3 0 Salinity 3.9 12.3 20 30.7 Filamentous algae none none none large amount Table 5: Zooplankton counts (number/m3 ) from night tows on 16/8/05 using a 100 µm mesh net. Tow duration was 2 minutes. Values at each site are an average of 3 tows plus/minus the standard error. Light and chlorophyll-a, Sanderson 4 28 Discussion Upstream chlorophyll-a concentrations are high whereas those near the mouth of the ocean are low, reflecting ocean conditions. The fact that chlorophyll-a drops rapidly as salinity increases shows that phytoplankton in the fresh water are lost as they are mixed into more saline water. Indeed, Aulacoseira counts decay exponentially with distance from the near-freshwater conditions in the upper estuary. Similarly, it seems that oceanic species, eg Thalassiosira, can be abundant in the lower estuary but have counts that are low upstream. Dinoflagellates and un-identified nanoplankton were much more abundant in mid-estuary than near either upstream or downstream boundaries — indicating that conditions within the estuary are favourable for these two groups. Dilution experiments showed high potential growth rates with low grazing rates in the estuary (salinities greater than 6 and less than 20). Further, the phytoplankton growth was not limited by nitrogen. Incubations in the laboratory are under saturating light conditions. They are also at slightly higher temperatures than the estuary. It is likely that the availability of light limits phytoplankton growth in the estuary. Benthic primary producers are obtain energy from the photosynthetically active radiation reaching the bottom. The fraction of surface light reaching the bottom is shown as a function of depth and light attenuation (κ) in Figure 16. Where the water column is deep benthic primary producers are unlikely to get sufficient light. Assume phytoplankton are vertically mixed through the water column on a time scale comparable to that over which they grow. Then the vertically-averaged light is an important factor for controlling primary production of phytoplankton. Figure 17 plots the ratio of vertically-averaged light to surface light as a function of both water depth and light attenuation κ. Clearly light attenuation and vertical mixing are going to be important factors controlling phytoplankton growth within the Hunter River Estuary. Hypsometry and an estimation of the vertically-averaged photosynthetically active radiation would be a first step for modelling primary production in this estuary. Hypsometry can be cal- Light and chlorophyll-a, Sanderson 29 2 −1 κ (m ) 1.5 0.1 1 0.5 1 2 3 4 Water Depth (m) 5 6 Figure 16: The ratio of bottom light to surface light is contoured as a function of water depth and light attenuation κ. The contour interval is 0.05. The dark contour is labelled. culated from bathymetric data. More sophisticated treatments should consider the vertical stratification using a vertical mixing model and the tendency for baroclinicity to stabilize the water column. Frequent measurement of the vertical and horizontal structure of chlorophyll would provide a data set for development of a primary production model. It must be stressed that the present chlorophyll-a measurements were of near-surface waters. A proper understanding requires measuring vertical profiles. Light and chlorophyll-a, Sanderson 30 2 0.2 −1 κ (m ) 1.5 0.4 1 0.5 1 2 3 4 Water Depth (m) 5 6 Figure 17: The ratio of vertically-averaged light to surface light is contoured as a function of water depth and light attenuation κ. The contour interval is 0.1. The dark contours are labelled.
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