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2012 SOUTHEASTERN NATURALIST 11(2):263–278
Algal Community Composition from Kaolin Recovery
Ponds Located in Middle Georgia
Joseph N. Dominy, Jr.1 and Kalina M. Manoylov1,*
Abstract - This is the first floristic and ecological evaluation of small pond systems
developed over different periods of time after seized kaolin mining operations. Assessment
of the total algal assemblage was used to infer environmental conditions of the
aquatic habitats and also provided information about the ecological health and integrity
of the aquatic ecosystems. The main objectives of this study were to document algal
community composition and discern the amount of time it takes for a mined pond to
reach its high biodiversity of primary producers. Winter and summer samples were taken
from a pond developing for two years after removal of kaolin and from a pond that was
thirty years old. Both pond systems contained rich algal communities predominantly
from Cyanobacteria, Bacillariophyta, and Chlorophyta; however, the 30-year pond had
higher Shannon-Wiener diversity, richness, and evenness values in both sampling seasons.
In winter, filamentous Zygnematales dominated algal communities in the 2-year
pond, while the 30-year pond community was dominated by diatoms. In both sites, the
most taxon- rich group was green algal representatives of Desmidiaceae. In summer,
potentially toxin- producing filamentous Cyanobacteria of the Nostocales were recorded
in the 2-year pond, while the 30-year pond had higher average algal cell evenness and
few toxic Nostocales. The average abundance of 11 diatom species, seven green algae,
and one representative each of Euglenophyta, Cyanobacteria, and Cryptophyta resulted
in less than 20 percent overall similarity between the two ponds. Our findings suggest
that after two years of development potentially harmful kaolin residues are removed by
natural sorption processes and do not negatively influence primary producers. However,
stabilization processes in those manmade ecosystems may potentially take more than two
years to produce high species richness and prevent toxic algal blooms.
In aquatic systems, diverse algal communities can be identified and used to
infer the overall health of the environment. Researchers have considered ecological
restoration or creation of new aquatic habitats as a means of increasing
ecological value of degraded lands and conserving biological diversity (Bradshaw
1983, Cairns and Heckman 1996, Hobbs and Norton 1996, Naveh 1994).
An example, of degraded lands, are those resulting from mining activities such
as kaolin production. Central Georgia is rich in kaolin. Kaolin, or alumina-silicate,
is a clay-like substance used in the production of rubber, plastics, paints,
paper, and many other products. Kaolin clay is extracted from many locations in
central Georgia and plays an important economic role in the state. The process of
kaolin mining takes several years and involves many stages in which the kaolin
!Department of Biological and Environmental Sciences, Georgia College and State University,
Milledgeville, GA 31061. *Corresponding author - email@example.com.
264 Southeastern Naturalist Vol. 11, No. 2
is removed. The process involves huge “cuts”, usually around 2 ha in size and
with varying depth, in which the kaolin is removed from the area. After mining
operations have ceased many sites are filled with water to produce man-made
ponds, which have unclear ecological functions. Kaolin companies are vested in
properly restoring these mined areas, where water quality is monitored regularly
to produce an environment desirable for aquatic organisms (NRC 1992). Afterwards,
the reclaimed lands are declared as suitable for a variety of uses including
agriculture, forestry, and wildlife use (Georgia Mining 2011).
Creation of aquatic ecosystems is deemed a good restoration practice (Brookes
and Shields 1996) with the potential of complete ecosystem structure and function.
The time it takes for ecosystem structure and function to reach their full
potential is unknown. Full ecosystem potential is measured with community attributes
like high biodiversity, high evenness, and absence of toxic algal blooms
(Chalar 2008). Biodiversity assessment can allow for accurate prediction of
ecosystem stability (McGrady-Steed et al. 1997). Maintenance of ecosystem integrity
and preservation of biological diversity translates into valuable outcomes
for any natural habitat and should not be measured with monetary or moral values
(Pimentel et al. 1997).
Ecosystem-wide research for manmade small lakes and kaolin recovery ponds
is limited. Biological research of the relationship between algae and kaolin residue
has been dominated by experimental work done by Guenther and Bozelli
(2004). Algae and kaolin particles have been used for understanding sedimentation
and sorption processes (Cuker 1987) as well as algal survival after kaolin dust
coverage. Inorganic particles, dissolved oxygen, total phosphorus, and iron were
reported to decrease over time in small recovery ponds, while other metals and
organic carbon increased with time (Kalin et al. 2001). Due to the high remnant
amounts of silicates and silicic acid, diatoms were reported as dominating clay
and kaolin recovery ponds in Austria (Schagerl et al. 2010). A man-made pond
over a landfill in Virginia was dominated by pennate diatoms and chlorophytes
together with common Cyanobacteria after 10 years, while the algal community
in the same pond was dominated by centric diatoms and coccoid greens after 20
years (Sheavly and Marshall 1989).
Ponds provide important habitats for diverse floral and faunal communities,
including a number of rare taxa of conservation interest (Wood et al. 2003). This
paper examines the biodiversity and community attributes of primary producers
from two recovery ponds after kaolin extraction. It was hypothesized that algal
biodiversity can be used to predict recovery time of ponds after kaolin operations.
Based on extensive literature search, this report is the first on algal species
composition from kaolin recovery ponds in middle Georgia.
Material and methods
Algal communities were studied from aquatic locations at Gibraltar Mine,
Wilkinson County, GA. Sampling sites were located within the Coastal plain
region. Originally, 2-month, 2-year, 15-year, and 30-year recovering ponds were
2012 J.N. Dominy, Jr. and K.M. Manoylov 265
sampled. Due to safety procedure and restricted access, samples were collected
during the months of February/March 2010 (winter samples) and June 2010
(summer samples). The sites were chosen based upon their age and repeated access.
No algae were observed in the 2-month pond, and the 15-year pond was
sampled only once due to limited access.
The 2-year pond was located at 32°53'8.55"N, 83°5'30.23"W and had a surface
area of about 0.53 ha with a maximum depth of approximately 2 m (Fig. 1;
Georgia Encylopedia 2011). Low buffer zones covered with sparse grasses along
embankments characterize the surrounding area. The 30-year pond was located at
32°54'15.70"N, 83°6'26.95"W and had a surface area of 3.4 ha with a maximum
depth of 4 m. Increased vegetation surrounded the area, along with a tree line
located 7 m upward from the embankment.
Six samples in triplicate were obtained from each site and analyzed. The six
distinct composite samples were taken using scrapings from various aquatic plants
for epiphytic and attached algae, sediments for benthic algae, and three throws of
a Turtox Tow Net (Wildco Ltd.) for water column collection for planktonic algae.
Figure 1. An overview picture of the State of Georgia, Wilkinson County (indicated by a
star), and the 30-year pond (top inset image) and the 2-year pond (bottom inset image).
266 Southeastern Naturalist Vol. 11, No. 2
Using composite samples decreases the amount of pseudoreplicates within a site
(Hurlbert 1984). Water temperature and pH levels were measured using a YSI
556 Multiprobe System (YSI Inc., Yellow Springs, OH) at the time of collection
(APHA 2005). Summer water samples for chemical analyses were collected in
500-ml acid-washed polyethylene bottles and sent to the University of Georgia for
chemical analysis of inorganic nutrients, hardness, and ionic composition (Analytical
Chemistry Laboratory, www.swpa.uga.edu). Water with hardness less than
85 mg/L is considered good quality drinkable water (APHA 2005).
All samples were brought to known volume for relative cell density estimation
(relative cells per mL) and concentrated. Fresh samples were kept on ice and
observed within 2 hours of collection for best possible taxonomic identification
using the natural color of algae and organelle structures. Algal samples for enumeration
were preserved within 5 hours of collection in formaldehyde (3 percent
final concentration). At least 300 natural algal units were enumerated in each
sample. Enumeration of preserved samples was done in Palmer-Maloney counting
chambers (Palmer and Maloney 1954). Natural counting units were defined
as one unit for each colony, filament, diatom frustule (regardless if colonial or
filamentous), or unicellular alga. If counting only individual cells, a large colony
might underrepresent the number of algal taxa within complex photosynthetic
communities; therefore, a minimum number of algal units were enumerated in
each sample. Cells within a colony or filament were recorded. When colonies
were multilayered and direct observation of individual cells was not possible,
cell numbers were based on closest estimation. Identification of species continued
after enumeration by scanning slides and recording presence of new algal
species until no new taxa were observed on two consecutive semi-permanent flat
slides. Species richness and cell relative abundance was calculated based on the
total documented species (fresh observation after collection, 300-unit counts,
and post-counts scanning). Observations for algal identification to the lowest
taxonomic level possible were done at 400X under a Jenalumar (AusJena) microscope
equipped with differential interference contrast (DIC), and images were
captured with a Leica DM2500 color digital camera. Additional observations
for taxonomic diversity were done on flat slides and under higher magnification
as necessary. Current taxonomy followed Wehr and Sheath (2003), and the following
references for specific algal groups were consulted: Bacillariophyceae
- Krammer and Lange-Bertalot (1986, 1988, 1991a, 1991b), Patrick and Reimer
(1966, 1975); Chlorophyta - Komárek and Fott (1983); Chrysophyceae - Starmach
(1985); and Cyanobacteria - Komárek and Anagnostidis (1999, 2005).
Collected material was archived and deposited as part of the algal collection of
the GCSU Natural History Museum.
For this study, dominant taxa were defined as more than 20 percent relative
abundance of both units and cells from total recorded. Color digital images were
taken of all taxa occurring in a sample at a relative abundance of five percent or
more, with a goal of representing all taxa encountered at abundance of two percent
or more in the study. Unknown taxa, potentially new to science were well
2012 J.N. Dominy, Jr. and K.M. Manoylov 267
Species richness, species evenness, and species diversity were calculated for
each pond for all sampling dates. Species richness (S) describes the number of
species (varieties or forms) recorded from a pond. The Shannon Index (H') considers
species richness documented and proportion of each species out of total
abundance recorded (Shannon and Weaver 1949). Maximum biodiversity (Hmax)
was calculated as the natural logarithm of the documented total species richness
number (Hmax = ln[S]). Species evenness (J'; Pielou 1969) was calculated as a
proportion of the Shannon diversity and maximum biodiversity documented using
the formula J' = H'/Hmax. Species diversity is a measure of ecosystem function
that increases with either species richness or species evenness.
Two community similarity indices that take under consideration presence/
absence of species and are not biased by small sample size or relative proportions
were calculated. The Sǿrensen (1957) similarity index was calculated as
S = 2C /(A + B), where C is the number of common species between two sites, A
is the number of species in one site, and B is the number of species in the other
site. Similarity varies from 0 to 1; high similarity is expected at 0.7 and above.
For each pond as a whole, community similarity was calculated with the Jaccard
index (Jaccard 1901), which is the number of species found at both locations (j)
divided by the number of unique species (r) to either location combined (JI = j/r
To test if physiochemical characteristics differed between the sites, each
characteristic was compared with a t-test (α = 0.05). If the assumptions for t-test
were not met (e.g., normality, equal variance) and if transformation did not help,
the Mann-Whitney non-parametric test was used. Descriptive statistics were conducted
to analyze environmental variability and relationships between variables.
Statistical analyses were performed with SYSTAT® 13 (Wilkinson 1989).
A one-way ANOSIM was used to compare the ponds based on species’ relative
abundance (untransformed and transformed data) with season nested within
pond type. ANOSIM calculates test statistics, the R statistics, and gives stress
values as a measure of the significance of the two-dimensional placement of sites
on the ordination (stress value of more than 0.2 is considered random, values
below are significant). ANOSIM global R evaluates whether there is a significant
difference in species composition between two or more groups of sites. Stress
values are goodness-of-fit statistics for NMDS.
Contribution to dissimilarity between the sites was presented as percentage
of contribution of each taxon to the overall dissimilarity (D) between the two
ponds of different ages as quantified by the SIMPER routine (Clarke and Warwick
2001). The procedure calculates the ratio of the mean dissimilarity for all
sample pairs between groups to standard deviation (SD) (mean D/SD[D]). This
ratio indicates how consistently a taxon contributes to mean dissimilarity across
all pairs and is a measure of the importance of each taxon in discriminating community
composition at each sampling date (Clarke and Warwick 2001).
268 Southeastern Naturalist Vol. 11, No. 2
Physiochemical properties between the two sites remained relatively constant
and were not significantly different, with the exception of winter and summer
average temperatures (Table 1). Water temperature varied in winter around 11 °C
in the 2-year and around 13 °C in the 30-year pond, while summer temperatures
in both ponds were 20 °C higher. Acidity levels in the 2-year pond were lower
(average pH 6.4) than in the 30-year pond (close to 7). All measured parameters
indicated low nutrient and ionic composition. Both ponds had low concentrations
of calcium and magnesium. Hardness remained below the standard (around 15
mg/L) in both ponds.
Numbers of algal units varied more for the 2-year pond (mean = 1383, standard
deviation = 621, range = 466–2328) compared to the 30-year pond (mean =
893, standard deviation = 216, range = 553–1124). Diverse species composition
was documented for both ponds. Representatives of seven algal divisions were
documented, and dominant algae belonged to Bacillariophyta or Chlorophyta.
Species richness was equal or lower in the 2-year pond, with highest richness
reported during summer in the 30-year pond (Fig. 2). A total of 81 species were
identified in the 2-year pond, and 132 species were identified in the 30-year pond.
The winter community differed in the two sites. The 2-year pond was dominated
by filamentous green algae (Hyalotheca dissiliens and Mougeotia sp.) and coccoid
algae, while the 30-year pond was dominated by chain-forming diatoms
and filamentous green algae. At the beginning of summer, algal community in
the 2-year pond was dominated by colonies of Tetraspora cf. cylindrica (Wahl)
Agardh, while no other algae reached 20 percent relative abundance later in the
Table 1. Physiochemical characteristics for the two locations, given mean ± S.E, all means not
significantly different (P > 0.05) between the 2 ponds.
Sample ID 2-year pond 30-year pond
Temp (°C) 10.9 ± 1.2 13.4 ± 1.9
pH 6.4 ± 0.5 7.2 ± 0.4
Temp (°C) 29.3 ± 0.17 30.3 ± 1.00
pH 6.4 ± 0.11 6.7 ± 0.10
NO2-N + NO3-N ( μg/L) 4.4 ± 0.9 5.3 ± 2.0
NH4-N ( μg/L) 46.2 ± 15.70 48.4 ± 0.65
PO4-P ( μg/L) 1.50 ± 0.3 2.01 ± 1.3
Ca (mg/L) 1.56 ± 0.0 7.50 ± 5.8
Mg (mg/L) 0.81 ± 0.08 1.40 ± 0.50
Hardness (mg/L) 2.4 ± 0.08 8.9 ± 6.30
K (mg/L) 0.71 ± 0.23 0.90 ± 0.10
Na (mg/L) 0.95 ± 0.19 1.01 ± 0.20
Fe ( μg/L) 13.6 ± 9.2 50.5 ± 27.6
Cl (mg/L) 2.26 ± 0.46 2.27 ± 0.50
SO4 (mg/L) 7.12 ± 1.1 7.20 ± 1.2
fl( μg/L) 114.6 ± 1.9 133.7 ± 16.6
2012 J.N. Dominy, Jr. and K.M. Manoylov 269
summer. At this time, the community was dominated by long filaments of green
algae and blue-green bacteria.
The rarest alga reported from the 2-year pond’s summer samples are potential
toxin-producing and bloom-forming blue-green bacteria (Fig. 3).
Morphologically, Cylindrospermopsis raciborskii (Woloszynka) Seenaya & Subba
Raju produced longer conical heterocytes and intercalary akinetes, sometimes
two in a row (Figs. 3a–d). This is the first report for Georgia of the heterocyteproducing
filament Richelia siamensis (Antarikanonda) Hindák (Fig. 3e), a
species with undetermined ecology. At least one new species to science, of the
genera Cylindrospermum, was documented in this study (Fig. 3f).
Within the 30-year pond, all algae counts were dominated by chains of
Fragilaria crotonensis Kitton and Fragilariforma sp. together with colonies
of Tetraspora gelatinisa (Vaucher) Desvaux. In the 30-year pond, green filamentous
algae such as species of Oedogonium and Desmidium swartzii Agardh,
were found in large quantities during the winter months, whereas in the summer
months, Cyanobacteria filaments like Oscillatoria princeps Vaucher and Nostoc
sp. were dominate. The diatoms like Encyonema silesiacum (Bleisch) Mann,
Craticula cuspidata (Kützing) Mann, Epithemia adnata (Kützing) Brébisson,
and Rhopalodia gibba (Ehrenberg) Müller were found in small abundances.
Filamentous green algae such as Spirogyra sp. and diverse coccoid members
of Desmidiaceae were reported with representative species of Micrasterias,
Closterium, Staurastrum, and Cosmarium present. A fairly rare green colony
Gloeotaenium loitelsbergerianum Hansgirg was also documented.
Overall, 21 species accounted for a total of 74.29 percent dissimilarity between
the two ponds, with six of those species contributing at least 3 percent of
the dissimilarity (Table 2). Twenty-eight percent (43 out of total 151 algal species
observed) of the taxa observed carried 90 percent dissimilarity of the ponds. Two
Figure 2. Species richness along sampling events: 2-year pond (striped bars) and 30-year
pond (solid bars). Numbers coincide with the sampling events that took place: 1–3 in
winter, 4–6 in summer.
270 Southeastern Naturalist Vol. 11, No. 2
species, Fragilaria crotonensis and Mougeotia sp. 2, contributed 12 percent and
9.3 percent, respectively, to the overall dissimilarity.
The Shannon-Wiener diversity values for the 2-year pond had an overall average
of 1.70, whereas the overall average diversity for the 30-year pond was 2.17.
On average, the diversity was highest in the 30-year pond and lowest in the winter
collection of the 2-year pond (Table 3). The 2-year pond community exhibited
only slight variations in their cell algal community structure (max. diversity =
2.83; min = 0.88), whereas the 30-year pond community was closer to the predicted
maximum biodiversity (Table 3). On average, the winter community was
Figure 3. Heterocyteforming
the 2-year recovery
pond: a–d. Cylindrospermopsis
Seenaya & SubbaRaju,
e. Richelia siamensis
Hindák, and f. Cylindrospermum
Scale bar equal to 10
2012 J.N. Dominy, Jr. and K.M. Manoylov 271
lower in biodiversity and higher in evenness. Positive significant correlation between
algal biodiversity and pond age was found for two of the summer sampling
events (Fig. 4)
Sampling date of specific community composition varied within the pond,
with two to 19 similar taxa between the two ponds (24 total common for the two
sites). This finding corresponded to lower average community similarity during
summer collection compared to winter (winter mean = 0.3, range = 0.16 to 0.43;
summer mean = 0.21, range = 0.10 to 0.34). The highest number of the same taxa
composition (19) was recorded in the site with the highest species richness, yet
similarity was only 34 percent. The lowest similarity reported was 10 percent for
the last summer collection (Table 4). Community similarity for the two ponds
was distinctly different based on total recorded algal composition and a low Jaccard
index of 19.5 percent.
Table 2. Average relative abundance (RA) of algal taxa that contributed more than one percent
to the overall dissimilarity between 2-year and 30-year ponds recovering after kaolin extraction.
The percent contribution (%D) of each taxon to overall dissimilarity (D) between the two ponds
is shown; D/SD(D) is the ratio of the mean D for all sample pairs between pond types to the
standard deviation of D and is a measure of the importance of each taxon in discriminating communities
between pond types. Contribution (Con) of each taxon is given also and the cumulative
percent (Cum%) is calculated.
Taxon Group 30-y 2-y %D D/SD(D) Con Cum%
Fragilaria crotonensis Kitton Diatoms 67.7 10.0 10.3 1.20 12.0 11.99
Mougeotia sp. 2 Green 2.8 48.8 8.0 0.98 9.3 21.31
Tetraspora gelatinosa (Vaucher) Desvaux Green 33.3 0.0 5.5 0.44 6.4 27.75
Hyalotheca dissiliens (Smith) Brébisson Green 2.8 27.0 4.5 0.74 5.2 32.99
Eunotia parallela Ehrenberg Diatoms 21.2 14.3 3.4 1.14 4.0 36.97
Fragilariforma sp. 1 Diatoms 18.3 0.0 3.0 0.44 3.5 40.50
Mougeotia sp. 1 Green 3.8 16.8 2.8 1.02 3.2 43.74
Tetraspora cf. cylindrica (Wahl) Agardh Green 0.0 16.3 2.7 0.44 3.2 46.92
Oedogonium sp. 2 Green 16.8 1.3 2.6 1.45 3.1 49.99
Navicula cryptotenella Lange-Bertalot Diatoms 2.3 17.3 2.6 1.01 3.0 53.03
Nitzschia dissipata (Kützing) Grunow Diatoms 0.8 16.3 2.6 1.18 3.0 56.00
Navicula trivialis var. oligotraphenta Diatoms 13.5 3.5 2.4 0.87 2.8 58.83
L-Bert. et Hofmann
Achnanthidium minutissimum (Kützing) Diatoms 6.0 16.5 2.3 1.31 2.6 61.46
Oedogonium sp. 1 Green 8.7 2.0 1.6 0.58 1.9 63.35
Surirella tenera Gregory Diatoms 0.7 9.7 1.6 0.97 1.9 65.21
Frustulia krammeri Lange-Bertalot et Diatoms 1.3 9.7 1.5 0.87 1.8 67.00
Trachelomonas sp.1 Euglenoid 8.2 0.3 1.4 0.47 1.6 68.59
Stenopterobia delicatissma (Lewis) Diatoms 0.0 8.0 1.3 0.44 1.6 70.15
Leptolyngbya sp. 1 Cyanophyta 5.2 9.7 1.3 1.3 1.5 71.61
Cryptomonas ovata Ehrenberg Cryptophyta 0.0 7.2 1.2 0.404 1.4 73.00
Synedra ulna (Nitzsch) Ehrenberg Diatoms 5.7 2.8 1.1 0.75 1.3 74.29
272 Southeastern Naturalist Vol. 11, No. 2
The ordination plot showed significant separation of the ponds based on
years of recovery after kaolin removal, and this separation was independent of
season (Fig. 5). The separation of ponds based on untransformed (ANOSIM
Table 3. Community indices of ponds with two years and 30 years of development over two seasons
with three replicas reported. H' = Shannon diversity, H' max = maximum Shannon diversity for each
site, and J' =Evenness for units and cells.
Pond Units Cells
age Season Replica H' H' max J' H' H' max J'
2 Winter 1 1.78 2.56 0.69 1.61 2.56 0.63
2 Winter 2 2.06 3.22 0.64 1.16 3.22 0.36
2 Winter 3 2.71 3.30 0.82 2.14 3.30 0.65
2 Summer 1 2.63 3.78 0.69 0.88 3.78 0.23
2 Summer 2 2.64 3.33 0.79 2.46 3.33 0.74
2 Summer 3 2.64 3.00 0.88 1.97 3.00 0.66
30 Winter 1 1.34 2.30 0.58 1.68 2.30 0.73
30 Winter 2 1.66 3.26 0.51 1.44 3.26 0.44
30 Winter 3 2.36 3.43 0.69 2.23 3.43 0.65
30 Summer 1 3.19 4.22 0.76 2.94 4.22 0.70
30 Summer 2 3.17 4.09 0.77 2.64 4.09 0.64
30 Summer 3 1.47 3.09 0.48 2.07 3.09 0.67
Figure 4. Regression analysis of Shannon diversity change for the 2-year and 30-year
2012 J.N. Dominy, Jr. and K.M. Manoylov 273
global R2 = 0.65, NMDS stress = 0.11,) and square root transformed data
(ANOSIM global R2 = 0.48, NMDS stress = 0.1) was significant. Separation was
based on presence-absence of taxa in one pond or another and not simply a result
of differences in relative abundance.
Biodiversity was lower for the 2-year pond, and the algal community experienced
potential turnover from filamentous green algae to filamentous blue-green
bacteria for both sites. Toxin-producing cyanobacteria, Cylindrospermopsis
rociborskii, and members of the genus Cylindrospermum were observed during
analysis only in the summer samples in the 2-year pond; however, the concentrations
found were too low for bloom formation. Biodiversity for the 30-year pond
remained significantly higher during both seasons. Findings of this study suggest
Figure 5. Nonmetric multidimensional scaling ordination plot of algal communities from
the 2-year and 30-year ponds in Georgia based on Bray-Curtis dissimilarity and presenceabsence
of square-root transformed data: winter (w) and summer (s).
Table 4. Sǿrensen Similarity (S) presented as percent similarity of the 2 ponds collected on the
same date. Numbers coincide with the sampling events that took place: 1–3 in winter, 4–6 in summer.
Season Sample Similar species S
Winter 1 5 0.43
Winter 2 4 0.16
Winter 3 9 0.31
Summer 4 19 0.34
Summer 5 9 0.20
Summer 6 2 0.10
274 Southeastern Naturalist Vol. 11, No. 2
that the 30-year pond can be used as an ecological benchmark for implications
referring to an ecological, stabilized aquatic system after mining processes have
ceased. Identification of high-value ecological benchmarks has been reported as a
high priority for aquatic science (Hawkins et al. 2010). A thirty-year time period
of development has been reported as a good benchmark of stability in terrestrial
communities (Baeten et al. 2010).
Diverse algal communities increase the stability of an area and therefore
the overall health of an ecosystem. Similar to other research on small ponds
(Sheavly and Marshal 1989), biodiversity of the ecosystem recovering from
kaolin extraction increased with time. Dominant green filamentous algae
from genera like Mougeotia, and Hyalotheca were observed during winter
months as large amounts of algal units and may suggest mesotrophic conditions.
Mougeotia can form substantial sub-surface growths in acidified waters and is
widely regarded as an indicator of early environmental change (Turner et al.
1991). In the 2-year pond, pH was consistently lower, but both ponds had very
diverse representativeness of Desmidiaceae. Other green filaments were represented
with few very long filaments and comprised a large portion of the cell
counts. Results from this study suggest that after two years of pond reclamation,
environmental conditions allowed for the development of a diverse algal
community. It was found that for both sites, as the overall water temperature
increased, there was an increase in algal diversity and abundance. This finding
was to be expected since green algae and Cyanobacteria prefer warmer conditions
and increased sunlight (Stevenson 1996). Abundant algae from the winter
sampling event in the 2-year pond were not observed in consecutive sampling
events, documenting dynamic changes in the algal community composition.
Our results are a snapshot of the algal community development under similar
environmental conditions in ponds of different age, and we are not addressing
short-term successional changes through time.
High diversity documented in this study is similar to other diverse algal
communities developed in low-nutrient clay pits in urban areas (Schagerl et al.
2010). The primary mechanism driving these relationships was a dominance
(or selection) effect: more diverse communities were more likely to contain the
most productive and least prone to invasion types of algae. Low evenness due
to dominant species can produce a negative relationship between species diversity
and ecosystem function (Creed et al. 2009). Regardless of the extensive
observation in the 30-year pond, summer toxin-producing Cylindrospermopsis
rociborskii representatives were not documented. The highest relative abundance
recorded in this study in 30-year summer samples (Fragilaria crotonensis chains
or Tetraspora colonies) both corresponded to different measures on the units and
cell diversity and evenness metric. Algal taxonomy and biology is an important
contributing factor for the diverse responses in kaolin recovery ponds.
Decrease in available nitrogen together with high water temperature (Nydick
et al. 2004) will lead to an increase in Cyanobacteria (Blue-green algae) with
heterocytes. This result was observed in the 2-year pond summer sampling.
Observations at the 2-year pond, where constant runoff of sediment was not
2012 J.N. Dominy, Jr. and K.M. Manoylov 275
buffered, might provide an explanation of increased turbidity and high availability
of clay particles. The 30-year site is surrounded by well-established meadow,
forested area, and no agricultural activities, so observation of the low nutrient
algal taxa was not surprising. There are no urban areas or agricultural activities
in the immediate surroundings of both ponds.
Analyses of nutrient content among sites were conducted during the summer
months. When comparisons were analyzed between both sites, nutrient availability
seems to have played an insignificant role in determination of species richness
between the two locations. Cyanobacteria possess nitrogen-fixing heterocytes,
which allow them to thrive in conditions when nitrogen levels are low and thus
unfavorable for green filamentous algae. Chemical analyses during the summer
months revealed that distinctions between the two sites’ abundance and diversity
values of the algal species were not affected by nutrient concentrations. Studies
performed on terrestrial plants when examining the effect on resource availability
has shown that an increase in nutrients will enhance biodiversity effects within
an area (Jonsson and Malmqvist 2003). However, it has also been recognized
that within algal communities species richness may be independent of nutrient
availability, and the effect of species richness on the temporal variability of community
was neutral only in nutrient-rich environments, while a stabilizing effect
of diversity was found in oligotrophic environments (Zhang and Zhang 2006).
Both kaolin recovery ponds were considered nutrient-poor and presented a good
opportunity to compare algal community alterations over short and long periods
of time. One possible explanation of the high diversity found within the 30-year
site could be correlated to the “niche complementarity effect” (Zhang and Zhang
2006). This effect explains how resource partitioning within a stabilized aquatic
system may overcome the results of temporal change in an environment and
would explain the higher community diversity and species richness found within
the 30-year site in comparison to the 2-year site, as well as the documented
appearance of potentially bloom-forming blue-green algae during the summer
months in the 2-year pond. Longer-term development of aquatic systems with
minimum anthropogenic influence can lead to system-wide stability even for
newly created aquatic habitats after destructive human practices.
We would like to thank BASF Chemical Company, McIntyre, GA for providing supervised
access to the sites. We would also like to thank Mike McEwen for his help with
field collections. This work was part of the first author’s Senior Undergraduate Capstone
project at the Department of Biological and Environmental Sciences at Georgia College
and State University. We are very grateful to the editing team at Southeastern Naturalist;
in addition, the manuscript was greatly improved thanks to detailed reviews by two
American Public Health Association (APHA). 2005. Standard methods for examination
of water and wastewater. American Public Health Association, Washington, DC.
276 Southeastern Naturalist Vol. 11, No. 2
Baeten, L., M. Hermy, S. Van Daele, and K. Verheyen. 2010. Unexpected understory
community development after 30 years in ancient and post-agricultural forests. Journal
of Ecology 98:1447–1453.
Bradshaw, A.D. 1983. The reconstruction of ecosystems. Journal of Applied Ecology
Brookes, A., and F.D. Shields, Jr. 1996. River Channel Restoration: Guiding Principles
for Sustainable Projects. Wiley, Chichester, UK.
Cairns, J., and J.R. Heckman. 1996. Restoration ecology: The state of emerging field.
Annual Review of Energy and Environment 21:167–189.
Chalar, G. 2008. The use of phytoplankton patterns of diversity for algal bloom management
Clarke, K.R., and R.M. Warwick. 2001. Change in Marine Communities: An Approach to
Statistical Analysis and Interpretation. 2nd Edition. PRIMER-E Ltd, Plymouth, UK.
Creed, R.P., R.P. Cherry, J.R. Pflaum, and C.J. Wood. 2009. Dominant species can produce
a negative relationship between species diversity and ecosystem function. Oikos
Cuker, B.E. 1987. Field experiments on the influences of suspended clay and P on the
plankton of a small lake. Limnology and Oceanography 35:840–847.
Georgia Encyclopedia. 2011. Available online at http://www.georgiaencyclopedia.org.
Accessed 2 January 2012.
Georgia Mining. 2011. Georgia’s Kaolin Industry. Available online at http://www.Georgiamining.
org. Accessed 9 October 2011.
Guenther, M., and R. Bozeli. 2004. Factors influencing alga-clay aggregation. Hydrobiologia
Hawkins, C.P., J.R. Olson, and R.A. Hill. 2010. The reference condition: Predicting
benchmarks for ecological and water–quality assessments. Journal of the North
American Benthological Society 29(1):312–343.
Hobbs, R.J., and D.A. Norton. 1996. Towards a conceptual framework for restoration
ecology. Restoration Ecology 4:93–110.
Hurlbert, S.H. 1984. Pseudoreplication and the design of ecological field experiments.
Ecological Monographs 54:187–211.
Jaccard, P. 1901. Étude comparative de la distribution florale dans une portion des Alpes
et des Jura. Bulletin de la Société Vaudoise des Sciences Naturelles 37:547–579.
Jonsson, M., and B. Malmqvist. 2003. Mechanisms behind positive diversity effects on
ecosystem functioning: Testing the facilitation and interference hypotheses. Oecologia
Kalin, M., Y. Cao, M. Smith, and M. Olaveson. 2001. Development of the phytoplankton
community in a pit-lake in relation to water quality changes. Water Research 35
Komárek, J., and K. Anagnostidis. 1999. Cyanoprokaryota 1. Teil: Chroococcales. In H.
Ettl, G. Gärtner, H. Heynig, and D. Mollenhauer (Eds.). Süsswasserflora von Mitteleuropa
19/1, Gustav Fischer Verlag, Jena-Stuttgart-Lübeck-Ulm, Germany. 548
Komárek, J., and K. Anagnostidis. 2005. Cyanoprokaryota 2. Teil/ 2nd Part: Oscillatoriales.
In B. Büdel, L. Krienitz, G. Gärtner, and M. Schagerl (Eds.). Süsswasserflora
von Mitteleuropa 19/2, Elsevier/Spektrum, Heidelberg, Germany. 759 pp.
Komárek, J., and B. Fott. 1983. Chlorophyceae (Grunalgen) Ordnung: Chlorococcales.
Das Phytoplankton des Siifiwassers. In Die Binnengewasser XVI, 7(1). Gustav Fisher
Verlag, Stuttgart, Germany. 1044 pp.
2012 J.N. Dominy, Jr. and K.M. Manoylov 277
Krammer, K., and H. Lange-Bertalot. 1986. Bacillariophyceae. 1. Teil: Naviculaceae. In
H. Ettl, G. Gärtner, H. Heynig, and D. Mollenhauer (Eds.). Süsswasserflora von Mitteleuropa.
2(1). Gustav Fisher Verlag, Jena, Germany. 876 pp.
Krammer, K., and H. Lange-Bertalot. 1988. Bacillariophyceae. 2. Teil: Bacillariaceae,
Epithemiaceae, Surirellaceae. In H. Ettl, G. Gärtner, H. Heynig, and D. Mollenhauer
(Eds.). Süsswasserflora von Mitteleuropa. 2(2). Gustav Fisher Verlag, Stuttgart, Germany.
Krammer, K., and H. Lange-Bertalot. 1991a. Bacillariophyceae. 3. Teil: Centrales, Fragilariaceae,
Eunotiaceae. In H. Ettl, G. Gärtner, H. Heynig, and D. Mollenhauer (Eds.).
Süsswasserflora von Mitteleuropa. 2(3). Gustav Fisher Verlag, Stuttgart, Germany.
Krammer, K., and H. Lange-Bertalot. 1991b. Bacillariophyceae. 4. Teil: Achnanthaceae.
Kritische Ergänzungen zu Navicula (Lineolatae) und Gomphonema. In H. Ettl, G.
Gärtner, H. Heynig, and D. Mollenhauer (Eds.). Süsswasserflora von Mitteleuropa.
2(4). Gustav Fisher Verlag, Stuttgart, Germany. 437 pp.
McGrady-Steed, J., P.M. Harris, and P.J. Morin. 1997. Biodiversity regulates ecosystem
predictability. Nature 390:162–165.
National Research Council (NRC). 1992. Restoration of aquatic ecosystems: Science,
technology, and public policy. National Academic Press, Washington, DC. 552 pp.
Naveh, Z. 1994. From biodiversity to ecodiversity: A landscape-ecology approach to
conservation and restoration. Restoration Ecology 2:180–189.
Nydick, K.R., B.M. Lafranconcos, J.S. Baron, and B.M. Johnson. 2004. Nitrogen
regulation of biomass, productivity, and composition in shallow mountain lakes,
Snowy Range, Wyoming, USA. Canadian Journal of Fisheries and Aquatic Science
Palmer, C.M., and T.E. Maloney. 1954. A new counting slide for nanoplankton. American
Society of Limnology and Oceanography, Special Publication 21. 6 pp.
Patrick, R., and C.W. Reimer. 1966. The Diatoms of the United States. Vol. 1. Monographs
of the Academy of Natural Sciences of Philadelphia 13. 688 pp.
Patrick, R. and C.W. Reimer. 1975. The Diatoms of the United States. Vol. 2. Monographs
of the Academy of Natural Sciences of Philadelphia 13. 213 pp.
Pielou, E.C. 1969. An Introduction to Mathematical Ecology. John Wiley and Sons, New
Pimental, D., C. Wilson, C. McCullum, R. Huang, P. Dwen, J. Flack, Q. Tran, T. Saltman,
and B. Cliff. 1997. Economics and environmental benefits of biodiversity. Bioscience
Schagerl, M., I. Bloch, D.G. Angleler, and C. Fesl. 2010. The use of urban clay-pit ponds
for human recreational assessment of impact on water quality and phytoplankton assemblage.
Environmental Monitoring Assessment 165:283–293.
Shannon, C.E., and W. Weaver. 1949. The Mathematical Theory of Communication.
University of Illinois Press, Urbana, IL.
Sheavly, S.B., and H.G. Marshall. 1989. Phytoplankton composition in borrow pit lake in
Virginia. Proceedings of the Biological Society of Washington 102 (1):272–279.
Sørensen, T. 1957. A method of establishing groups of equal amplitude in plant sociology
based on similarity of species and its application to analyses of the vegetation on
Danish commons. Biologiske Skrifter / Kongelige Danske Videnskabernes Selskab 5
Starmach, K. 1985. Chrysophyceae und Haptophyceae. In H. Ettl, J. Gerloff, and D. Mollenhauer
(Eds.). Su ̈ßwasserflora von Mitteleuropa. Gustav Fischer Verlag, Stuttgart,
Germany. 515 pp.
278 Southeastern Naturalist Vol. 11, No. 2
Stevenson, R.J. 1996. An introduction to algal ecology in freshwater benthic habitat.
Pp. 3–30, In R.J. Stevenson, M.L. Bothwell, and R.L. Lowe (Eds.). Algal Ecology.
Academic Press, San Diego, CA.
Turner, M.A., E.T., Howell, M. Summerby, R.H. Hessleim, D.L. Findlay, and M.B. Jackson.
1991. Changes in epilithon and epiphyton associated with experimental acidification
of a lake to pH 5. Limnology and Oceanography 36:1390–1405.
Wehr, J.D., and R.G. Sheath. 2003. Freshwater algae of North America. Ecology and
Classification. Elsevier Science (USA), Academic Press, San Diego, CA. 918 pp.
Wilkinson, L. 1989. Systat: The System for Statistics. Evanston, IL. 822 pp.
Wood, P.J., M.T. Greenwood, and M.D. Agnew. 2003. Pond biodiversity and habitat loss
in the UK. Area 35(2):206–216
Zhang, Q., and D. Zhang. 2006. Resource availability and biodiversity effects of the
productivity, temporal variability, and resistance of experimental algal communities.