Tracking Recovery Patterns in Acidified Lakes: A Paleolimnological Perspective

Long-term data are often lacking to effectively assess patterns of lake acidification and recovery. Fortunately, paleolimnological techniques can be used to infer past changes in lakewater acidity and related variables by means of biological indicators, such as diatom valves and chrysophyte scales, preserved in ²¹⁰Pb-dated sediment cores. We summarize paleolimnological data that we have gathered from 36 Sudbury (Ontario) and 20 Adirondack Park (New York) lakes to estimate the magnitude of lake acidification and any subsequent recovery in these lake systems. In both regions, many lakes were shown to have acidified considerably, some over two pH units, since the 1850s. Although some recovery was noted in both lake regions, Sudbury lakes generally showed larger increases in inferred lakewater pH with recent declines in sulfur emissions. Possible explanations of these differences include the greater decrease in sulfate deposition in the Sudbury area, as well as generally longer residence times of lakes in Sudbury, perhaps allowing for more in-lake alkalinity generation. In addition, Sudbury lakes generally had higher pre-industrial pH levels, suggesting that lakes with higher natural buffering capacities are more likely to recover more quickly with declines in deposition, even if they had been acidified to a great extent.     

Paleolimnological investigation of three manipulated lakes from Sudbury,Canada

Stratigraphic changes in diatoms and chrysophytes from three manipulated Sudbury lakes were explored in an attempt to examine the influence of fertilization and/or neutralization on algal microfossil assemblages. Both diatom- and chrysophyte-inferred pH profiles indicate that the pH of Labelle Lake has remained fairly stable in the past. The study of Labelle and Middle lakes indicates that the addition of nutrients to acidic and non-acidic oligotrophic lakes did not directly influence diatom and chrysophyte species composition, perhaps because pH remained stable. The diatom and chrysophyte assemblages of Middle Lake only changed when the pH was raised. In Mountaintop Lake the recent shift in chrysophyte species composition and the resulting inferred pH decline is most likely related to a decline in mid-summer epilimnetic pH. Reliable paleolimnological inferences are difficult in lakes such as these because it is difficult to track limnological conditions in the absence of modern analogues.     

Elevated metal concentrations inhibit biological recovery of Cladocera in previously acidified boreal lakes

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Chrysophyte scales recorded in lake sediments from eastern Finland

Chrysophyte (mallomonadacean) scales were examined from the surface sediments of 18 lakes from eastern Finland, representing a wide range of limnological characteristics. Scales were always well preserved and 23 taxa were observed. The relative frequencies of the dominant taxa were plotted relative to the average autumn lakewater pH. The distribution of chrysophyte species in the calibration set was ordinated by means of DECORANA analysis. The occurrence of these species appeared to be closely related to pH, colour and nutrients in the ordinated lake groups. Chrysophyte scales are potentially useful environmental and paleolimnological indicators in eastern Finland.     

Geochemical controls for Al, Fe, Mn, Cd, Cu, Pb, and Zn during experimental acidification and recovery of Little Rock Lake, WI, USA

Concentrations of Al, Fe, Mn, Cd, Cu, Pb, and Zn were measured in thereference and treatment basins of Little Rock Lake (Vilas County, Wisconsin), alow-alkalinity, seepage system (pH 6.1, alkalinity25eq/L) during six years of a whole-basinacidificationand the first four years of the lake's recovery. The treatment basin wasacidified with H₂SO₄ in three two-year steps to pH5.6, 5.1, and 4.7. By the end of year 4 of recovery, treatmentbasin pH increased to 5.3 as a result of internal alkalinity generation.During acidification, dissolved Mn and Fe (0.4mpore-size filters) increased at pH 5.6; dissolved Al, Cd, and Zn becameelevated at pH 5.1; and dissolved Pb at pH 4.7. Dissolved Cu remainedsimilar in both basins to pH 4.7. Al, Fe and Mn levels declinedsignificantly during the recovery period, approaching values at pH 5.3intermediate between the concentrations at pH 5.6 and 5.1 during acidification.Dissolved Al and Fe in the reference basin were near the equilibrium levels forsolubility of gibbsite (Al(OH)₃) and amorphousFe(OH)₃(s).The acidified basin was undersaturated relative to gibbsite, and dissolved Alwas limited by pH disequilibrium between the water column and sediments andpossibly by Al-DOC precipitation. Dissolved Fe apparently was controlled bysolubility of amorphous Fe(OH)₃(s) and Fe-DOC precipitation.Dissolved Mn levels in both basins were consistent with manganite[-MnOOH(s)] solubility. Elevated levels of Cd, Pb, and Zn in thetreatment basin during acidification probably resulted from less efficientscavenging of atmospherically-deposited Cd, Pb, and Zn by settling particles.     

The effects of acidification on metal budgets of lakes and catchments

Metal (Cu, Ni, Zn, Fe, Mn and Al) budgets were measured for 5 lakes and their catchments near Sudbury, Ontario, an area severely affected by the emission and deposition of strong acids (H₂SO₄/SO₂) and metals. Three of the lakes were circum-neutral (pH 6.3–7.1) during the study period, while one lake had a pH of 4.8 and a fifth had very low pH ( 4.4).The lakes' catchments were all sources of Al, Mn and Ni, but were sinks for Cu and Zn. The Fe results were inconsistent; two lakes' catchments were sources while three were sinks.The acidic lakes were conservative (i.e. net retention of zero) with respect to Cu and Ni, while the circum-neutral lakes were effective sinks for these 2 metals. All of the lakes were sinks for Zn and Al, but the acidic lakes were less effective. All lakes were also Fe sinks. While there was no pattern relative to the lakes' pH's, there was a trend towards increasing Fe retention with increasing water replenishment time. The most acidic lake was actually a source of Mn, while the others were sinks.     

Immunoreactivity of the corpuscles of Stannius of the garpike,Lepisosteus osseus,to antisera against salmon and trout stanniocalcins

Stanniocalcin-immunoreactive cells were localized in the corpuscles of Stannius of a holostean fish, the garpike (Lepisosteus osseus), using antisera against salmon and trout stanniocalcins and the peroxidase-antiperoxidase and protein A-gold immunohistochemical methods. The stanniocalcin-immunoreactive cells were periodic acid-Schiff-positive, and antibody staining was abolished if the antiserum was preabsorbed with corpuscle homogenate. Immunocytochemistry revealed two reactive cell types in the glandular parenchyma, and immunoreactivity was confined to the secretory granules. Staining of the granules was also abolished when the antisera were blocked with crude corpuscle homogenate. When corpuscle extracts from garpike were subjected to sodium dodecyl sulphate-polyacrylamide gel electrophoresis and Western blot analysis, a single dense band was evident with a molecular weight of 68 kDa under non-reducing conditions, whereas three bands were observed (29, 31, and 34 kDa) under reducing conditions. Staining of all bands disappeared following preabsorption of the antiserum with salmon stanniocalcin, trout stanniocalcin, or garpike corpuscle extract. The results are compared with stanniocalcins from another extant holostean, the bowfin (Amia calva), and from more modern bony fishes, the teleosts.     

Cladoceran remains as indicators of lake acidification

Wide gaps in the records of limnological parameters from the past 30 to 50 years prevent a documentation of lake acidification. To solve this problem, methods must be found to infer former pH-conditions from sub-fossils. Therefore cladoceran remains were studied in undisturbed sediment cores of selected, poorly buffered lakes in the F.R.G. As a result, a correlation was found between the lakewater pH and the species composition of the assemblage of chydorid cladocerans. This correlation is highly significant and points out the distinct influence of low pH values (< 7) on this group of organisms. Reconstruction of the pH-history of three studied lakes evidences that acidification of poorly buffered lakes in the F.R.G. has taken place, even though it was not noticed previously.     

Regulation of bacterioplankton production and biomass in an oligotrophic cleanvater lake--the importance of the phytoplankton community

Estimations of bacterioplankton production and biomass werecarried out in enclosure experiments during two consecutiveyears (1989 and 1990) in oligotrophic clearwater Lake Njupfatet.The lake was limed in November 1989, and the experiments werecarried out both in 1989 (unlimed) and in 1990 (limed). Bags(3001) were manipulated with inorganic phosphorus and nitrogen,organic carbon, and metazoan zooplankton abundance. Both years,bacterial production was stimulated by inorganic nutrients aloneand in combination with organic carbon. However, the increasein bacterial production when inorganic nutrients were addedalone was much stronger in 1990 than in 1989. In 1989. bacterialproduction increased strongly only when inorganic nutrientsand organic carbon were added together. The phytoplankton communitywas dominated by the cyanobacterium Merismopedia tenuis-simaduring 1989, and the phytoplankton biomass increased only slightlywhen receiving inorganic nutrients. In 1990, when the lake hadbeen limed. M.tenuissima had completely disappeared and thephytoplankton community, dominated by Chrysophyceae and Chlorophyceae,responded strongly to additions of inorganic nutrients. Theincreased phytoplankton productivity in 1990 may have resultedin increased release of organic carbon, and this in turn thatthe carbon limitation of bacterioplankton production decreasedfrom 1989 to 1990. Zooplankton had a positive effect on bacterioplanktonproduction in 1989, but no effect in 1990. The loss of bacterialbiomass approximated 60% of the bacterial production in 1989,while in 1990 it almost equalled the bacterioplankton production.     

Effects of taurine on the phosphorylation of specific proteins in subcellular fractions of the rat retina

The effects of 20 mM taurine on the phosphorylation of specific proteins in mitochondrial and rod outer segment subcellular fractions of the rat retina were measured. A band of protein with an apparent molecular wieght of 20K was consistently inhibited by taurine. Densitometry measurements performed on gel electrophoresis autoradiograms from the mitochondrial fraction demonstrated a 42.7±8.3% decrease due to taurine (20 mM) in the area corresponding to radioactivity from the 20K phosphoprotein. However, only a 21.2±9.0% decrease was observed due to taurine in the rod outer segment preparation. These data suggest that taurine is exerting its primary effect on the phosphorylation of the 20K molecular weight protein in the mitochondria of the retina. In addition, calmodulin and phorbol ester had no effect on the phosphorylation of the 20K molecular weight protein.     

Effects of Changes in Acidity on Aquatic Insects in Rocky Littoral Habitats of Lakes Near Sudbur y, Ontario

Benthic aquatic insects were collected from rocky nearshore areas (< 1 m deep) of 17 lakes near Sudbury, Ontario, with a pH range of 4.7–7.3 and a size range of less than 10 ha to over 10,000 ha. These insect communities were composed of taxa common to lake soft-sediments and streams. Direct and indirect effects of lake acidity appeared to be major controls on the structure of these communities, implying that several factors may be involved in restructuring during acidification or recovery. Declines in abundances of several taxa of Ephemeroptera at pH below 5.5 were attributable to acid toxicity, while increases in the abundances of Odonata and Diptera at pH below 5.5 were associated with the absence of fish predators and other indirect effects of acidity. The communities of two experimentally neutralized lakes restructured rapidly within 5 years, approaching but not achieving community structures typical of our near-neutral survey lakes. Neutralization led to recolonization or increased abundance of the acid-sensitive mayfly, Stenacron interpunctatum , and the dragonfly, Boyeria grafiana ; however, recolonization by other taxa expected to be present in near-neutral lakes ( Stenonema femoratum, Eurylophella, and Basiaeshna janata ) was not observed. Consistent with results for the survey lakes, declines in the abundances of the dragonflies Aeshna interrupta, Aeshna eremita , and Leucorrhinia glacialis in the neutralized lakes were associated with reintroductions of Salvelinus fontinalis (aurora trout) and increased fish predation pressure, while reduced abundances of the dipterans Ceratopogonidae, Psectrocladius , and Stackelbergina may be related to indirect effects of acidity other than fish predation. Although community composition varied greatly across the acidity gradient, total species richness and abundance were not correlated with lake chemistry or number of fish species.     

Phytoplankton successions under ice cover in four lakes located in North-Eastern Sweden: Effects of liming

Phytoplankton successions under ice cover (January–March) were determined in four oligotrophic lakes (Burtjärn, Aspen, Vialamptjärn and Storkorstjärn) located in North-Eastern Sweden. The total phosphorus concentration in the lakes was less than 10 µg/L. Lake Burtjärn (reference lake) had a similar hydrology as Lake Aspen. Storkorstjärn and Vialamptjärn were of similar hydrology and had heavily colored water (>100 mg_Pt/L). Aspen as well as Vialamptjärn became continuously limed with calcium carbonate annually during the last decades. Biodiversity was considerably higher in the limed lakes (Aspen and Vialamptjärn) than in the untreated lakes (Burtjärn and Storkorstjärn). In Lake Burtjärn the most frequent species wereRhodomonas lacustris, Tabellaria flocculosa andBotryococcus braunii. Cryptophyceae (R. lacustris andCryptomonas marssonii) andDinophyceae (especiallyGymnodinium lantzschii) were common phytoplankton groups in Lake Aspen.Tabellaria flocculosa was also the most common organism in both humic lakes Storkorstjärn and Vialamptjärn, other phytoplankton groups were in the humic lakes scarce. Liming was found to have profound effects on phytoplankton communities studied.     

Lake trout demographics in relation to burbot and coregonine populations in the Algonquin Highlands,Ontario

The objective of the study was to test the hypothesis that lake trout populations change in relation to cisco, lake whitefish, round whitefish and burbot populations in lakes in the Algonquin Highlands region of Ontario. Lake trout population change is greatest where cisco and lake whitefish are present. Lake trout populations in lakes without either coregonine tend to have small adults and many juveniles. Where cisco or lake whitefish are present, adult lake trout are large, juvenile abundance is low, and the stock-recruit relationship appears to be uncoupled likely due to a larval bottleneck. Lake trout populations in these lakes may be sensitive to overfishing and recruitment failure. Lake trout populations do not appear to change in relation to round whitefish. There appears to be an indirect positive change on juvenile lake trout abundance through reductions in the density of benthic coregonines in the presence of large, hypolimnetic burbot.     

Paleolimnological evaluation of historical trophic state conditions in hypereutrophic Lake Thonotosassa, Florida, USA

We used paleolimnological methods to evaluate historical water quality in Lake Thonotosassa, Hillsborough County, Florida, USA. Sediment mapping shows that organic deposits are unevenly distributed in the lake. Two short (<130 cm) sediment cores from the depositional zone were analyzed for radioisotopes (²¹⁰Pb, ²²⁶Ra, and ¹³⁷Cs), bulk density, organic matter concentration, nutrients (C,N,P), and diatoms. ²¹⁰Pb results indicate that the profiles represent > 100 years of sediment accumulation. There is an abrupt change in sediment composition at about the turn of the century (80 cm depth), above which bulk density decreases and concentrations of organic matter, total C, total N, total P, and ²²⁶Ra activity increase. Diatom-based reconstructions of historical water-column trophic conditions indicate progressive nutrient enrichment in the lake during the past 100 years. Stratigraphic changes in diatom assemblages suggest that anthropogenic nutrient loading converted Lake Thonotosassa from a naturally eutrophic system to a hypereutrophic waterbody after 1900. Given the edaphic setting of Lake Thonotosassa, efforts to mitigate recent anthropogenic impacts will, at best, yield the eutrophic conditions that characterized the lake prior to human disturbance. This study illustrates the importance of paleolimnological data for targeting realistic water quality conditions when lake restoration is contemplated.Journal Series No. R-05019 of the Florida Agricultural Experiment Station     

Food limited survival of larval razorback sucker,Xyrauchen texanus,in the laboratory

Synopsis Razorback suckers in Lake Mohave, Arizona-Nevada, are suffering recruitment failure. Concomitantly, lake zooplankton levels are low and variable. We test, under laboratory conditions, the possibility that starvation is a cause of low recruitment. Razorback sucker larvae that were starved, received food too late, or were provided with insufficient food died between 20 and 30 d after hatching. Yolk absorption was at 8d after hatching, the critical period during which larvae must feed or most will die lies between 8 and 19d, and the point of irreversible starvation for individuals is between 19 and 23 d after hatching. Results support food-related mortality as a contributor to year-class failure of razorback sucker in Lake Mohave.     

A landscape approach to examining spatial patterns of limnological variables and long-term environmental change in a southern Canadian lake district

1. We explored patterns of limnological variables (physical, chemical and biological) with relation to landscape position (expressed as lake order) in 86 study lakes located on shield bedrock in south‐central Ontario, Canada. 2. Using anova s with lake order as the categorical variable, landscape position explained significant amounts of variation in major ion chemistry, physical and catchment characteristics, hypolimnetic oxygen, and community composition in algal (diatom, chrysophyte) and invertebrate (chironomid) assemblages preserved in surficial sediments. Several nutrient variables (TP, total phosphorus and TN, total nitrogen) and dissolved organic carbon did not have significant relationships with lake order. 3. The strongest relationships with lake order (as a fraction of variation explained in anova s) included silica concentrations (r² = 0.40) and SO₄ (r² = 0.29) concentrations, surface area (r² = 0.50) and hypolimnetic oxygen (r² = 0.29). 4. Bedrock geology (carbonate metasedimentary versus non‐carbonate bedrock) had strong influences on spatial gradients of pH and major ion chemistry. It was difficult to separate geological influences from spatial influences on limnological variables in this study, as drainage patterns in the region are highly influenced by surface features of underlying geological formations because of the very thin glacial till or exposed bedrock that exists in most catchments. 5. Patterns of limnological variables indicated that low‐order, headwater lakes had the lowest concentrations of major ions, and, from algal inferences of pH change, had been most susceptible to acidic deposition. High‐order, downstream lakes were larger and deeper, and had higher concentrations of hypolimnetic oxygen, indicating that optimal lake trout habitat was primarily located in high‐order lakes. 6. Variance partitioning analyses indicated that lake order as a metric of landscape position explained comparable portions of community variation in algal and invertebrate assemblages compared with geographic position (latitude, longitude) and Cartesian coordinate position (e.g. x, y, x², y², etc.) metrics. Lake order explained more community variation in chironomid assemblages compared with other landscape metrics, possibly because of the strong relationships between lake order and lake morphometry variables.     

Cost of predation avoidance in young-of-year lake trout (Salvelinus namaycush): growth differential in sub-optimal habitats

Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering     

SCALED CHRYSOPHYTES (CHRYSOPHYCEAE AND SYNUROPHYCEAE) FROM ADIRONDACK DRAINAGE LAKES AND THEIR RELATIONSHIP TO ENVIRONMENTAL VARIABLES1

The relationships between 23 scaled chrysophyte taxa (Chrysophyceae and Synurophyceae) and measured limnological variables in 62 Adirondack, New York, drainage lakes were examined by canonical correspondence analysis (CCA). The major proportion of variation in chrysophyte species distributions was strongly related to total monomeric Al (Al_m) and Mg concentrations, and their close correlates pH, Na, Ca, and acid-neutralizing capacity (ANC). Total monomeric Al concentrations explain a greater proportion of species variation than pH, suggesting that Al_m concentrations may be more important in governing the distribution of chrysophyte taxa in these lakes. Gaussian logit (GL) and linear logit (LL) regressions of the relative percentages of individual chrysophyte taxa to lakewater pH and Al_m concentrations and the examination of pH–Al_m response surfaces show that many chrysophyte taxa exhibit unique responses to these environmental gradients; taxa can be characterized as alkaline, circumneutral, acidic, and pH indifferent. Within each of these groups, taxa can be characterized further based upon their optima and tolerances to Al_m concentrations. Chrysophyte indicator species (i.e. a taxon with a strong statistical relationship to the environmental variable of interest, a well-defined optimum, and a narrow tolerance to the variable of interest) for pH include Mallomonas hindonii, M. crassisquama, M. pseudocoronata, and Synura uvella; M. hindonii, M. crassisquama, M. pseudocoronata, S. petersenii, and S. spinosa are good indicators of Al_m concentrations. Highly significant predictive models were developed to infer lakewater pH and Al_m concentrations from the relative percentages of chrysophyte scales in the study lakes. Model evaluation was based on their correlation coefficients and the root-mean-squared error of prediction (RMSE) derived from bootstrapping. Weighted averaging regression and calibration with tolerance down-weighting (i.e. weighting taxa inversely to their variance) produced superior results when compared to the computationally and data-demanding maximum likelihood methods and to simple weighted averaging regression and calibration.     

Contrasts between dystrophic and clearwater lakes in the long‐term effects of acidification on cladoceran assemblages

1. Most studies on zooplankton responses to acidification have focused on clearwater lakes with a dramatic acidification history. The role of dissolved organic carbon (DOC) in moderating zooplankton responses to acidification in naturally acidic, dystrophic lakes is less well understood and is partially impeded by a lack of baseline data. 2. Cladocera leave identifiable remains preserved in lake sediments that can be used to provide information on pre‐industrial species assemblages and their responses to environmental stressors such as acidification. Therefore, we used palaeolimnological approaches to track cladoceran assemblage responses to acidification since c.1850 (inferred from sedimentary diatom assemblages) in three acidified lakes in Kejimkujik National Park (Nova Scotia, Canada) that differ markedly in DOC content. These include two highly dystrophic lakes (Kejimkujik and Pebbleogittch lakes), and one clearwater lake (Beaverskin Lake). 3. In dystrophic Pebbleogittch Lake, an increase in the acid‐tolerant, jelly‐clad, pelagic taxon Holopedium glacialis occurred coincident with diatom‐inferred pH (DI‐pH) declines, but no other notable cladoceran assemblage shifts occurred. Similarly, Cladocera assemblages did not appear to respond to lakewater acidification in dystrophic Kejimkujik Lake. 4. In contrast, in the clearwater Beaverskin Lake, several observed shifts in cladoceran assemblage corresponded to DI‐pH declines, including an increase in the proportion of littoral taxa and an increase in Hill’s N2 species diversity. This may indicate increased water clarity as a result of acidification‐related decreases in DOC, which may have enhanced growth of emergent aquatic macrophytes and improved visibility for planktivorous fish, leading to increased predation on pelagic taxa. Species shifts within the littoral assemblage of Beaverskin Lake may reflect the differing tolerances of littoral taxa to low pH and aluminium toxicity. 5. Overall, our results suggest that cladoceran assemblages in naturally acidic, dystrophic lakes may be resilient against additional pH declines related to industrial emissions of acidifying agents, as dystrophic lakes are less vulnerable to increased aluminium toxicity and acidification‐induced increases in water clarity and often have a pre‐industrial cladoceran assemblage already adapted to acidic conditions.     

Life history,population dynamics and production of Pontoporeia hoyi (Crustacea,Amphipoda) in relation to the trophic gradient of Lake George,New York

The life history characteristics, population dynamics and production of Pontoporeia hoyi in Lake George, New York, were studied from May 1981 through October 1982. P. hoyi, in terms of both density and standing crop, is the most prevalent member of the deep water macrobenthos of Lake George. It reproduces in the winter, with young being released in the late winter-early spring. At the southernmost study site, young released in the spring grew to 6–7 mm in length and bred during their first winter. At the remaining sites, P. hoyi required two years to complete its life cycle. This difference in life history characteristics can be related to food availability and temperature differences. The open waters of the south end of Lake George are not only more productive but are also more closely associated with the littoral zone, providing a wealth of bacteria-rich detritus for benthic deposit feeders. The greater food availability in the south basin of Lake George is reflected in significantly larger brood sizes and smaller size at maturity for P. hoyi populations from the south end of the lake.The southernmost study site has significantly greater P. hoyi density and standing crop than all other sites. The cohort of the year dominated density and standing crop at the southern site while the cohort of the previous year dominated standing crop at the other sites. Peak abundance ranged from 600 · m^–2 at the north site to 2 900 · m^–2 at the south site. Cohort production ranged from 2g · m^–2 at the north site to 15g · m^–2 at the south site.