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A Study Comparing Ant Richness and Diversity Between Two Agricultural
Ecosystems: Polyculture versus Monoculture
Abagail E. Davis and Valerie S. Banschbach (Advisor)
Department of Environmental Studies, Roanoke College Salem, Virginia
29 April 2015
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ABSTRACT
In today’s agriculture, there are two types of farming methods: monoculture
and polyculture. Monoculture is born out of the industrial system and is when one
crop is planted on a single plot of land (Vandermeer, 2011). In contrast, polyculture
is typically smaller operations, organic, and the practice of planting a variety of
crops on a single plot of land (Vandermeer, 2011). Both agricultural methods
produce differing ecosystems. Typically found on large monoculture operations are
transgenic organisms like Bt-Corn. Corn is a crucial cash crop for small-scale and
large, industrial farmers with the United States leading the way in production and
exportation (Barton and Clark, 2014). However, with global climate change
constantly looming, farmers have observed an increase in corn pest species like the
European corn borer and corn earworm (Dively and Rose, 2003).
Therefore, agricultural researchers have developed a genetically engineered
corn crop composed of Bacillus thuringiensus, a gram-positive soil bacterium that
has insecticidal characteristics to combat pest issues on large-scale monocultures
(Sanchis and Bourguet, 2008). This study explored the question of whether differing
agricultural practices of monoculture and polyculture affect species richness and
diversity of ants on two farms: an polyculture farm called Lick Run Farm, and a
monoculture farm with transgenic corn called Michael Beahm Farm. Results indicate
species richness and diversity is higher on Lick Run Farm than on Michael Beahm
Farm. Higher diversity and richness is likely based on the monoculture At Michael
Beahm Farm versus polyculture agricultural methods at Lick Run Farm.
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INTRODUCTION
In an age of agricultural revitalization in the United States, consumers are
beginning to put more thought into their food purchases, and gravitating more
towards food items that have been cultivated sustainably (Irving et al., 2001). The
current agricultural paradigm in the United States is fueled by the industrial system
with goals of growth, profit, and productivity. This paradigm is driven by
monoculture, or the cultivation of a single crop on a plot of land (Vandermeer,
2011). Monoculture is favored in the industrial agricultural ecosystem because it is
less work than its alternative, and it speaks to productivity (Vaandermeer, 2011).
An alternative to monoculture; on the other hand, is polyculture. Polyculture is the
cultivation of many varieties of crops on a single plot of land together at the same
time (Vandermeer, 2011). Monoculture and polyculture, in essence, are differing
habitats. The biology and ecology for both forms of agriculture are differing. This
difference will be illustrated using the two farm sites used for this study: Lick Run
Farm and Michael Beahm Farm.
Lick Run Farm is an organic, but not certified, 3-acre polyculture farm in the
heart of the city of Roanoke, Virginia. It is surrounded by urban environment with
houses in every direction. Prior to becoming Lick Run Farm, the property was home
to a nursery called Crowell Nursery (Crawford, 2011). The soil prior to Lick Run
Farm ownership was degraded and lacking important nutrients. In an effort to
restore the soil, Rick Williams, the owner of Lick Run Farm, began planting cover
crops. Organic, polyculture farmers often use cover crops, or crops planted to add
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nutrients back to the soil and protect against erosion. Examples of cover crops are
legumes, various grasses, and grains (Zhang et al., 2007).
Williams also adds mulch and compost to his soil for extra enrichment.
Williams utilizes plant fragrances for pest management and to attract important
pollinators as well. Another important regime Williams uses on Lick Run Farm is the
three sisters method. The three sisters method utilizes intercropping of maize,
beans, and squash for efficient nutrient sequestration and uptake by the soil and the
plant, while also deterring pests. For instance, what nitrogen is taken up by the corn
the beans replenish back into the soil (Postma and Lynch, 2012).
Michael Beahm Farm is a 200-230 acre, monoculture farm in rural Hollins,
Virginia with 50-acres of cropland growing transgenic sweet Bt-corn. The Bt-corn
was not sprayed with any herbicides or pesticides. Michael Beahm Farm has been in
the Beahm family for several years now, and has always been operating as a larger-
scale monoculture operation.
Corn is a crucial agricultural commodity to the United States comprising
nearly 94 million acres of farmland in 2013. It is the most economically valuable
crop estimating a total production value of $67 billion (Barton and Clark, 2014).
Additionally, the United States is the number one producer and exporter of corn
comprising 36% of the global output. Unfortunately, natural events out of
humankind’s control are posing serious threats to the success of corn in the United
States. In 2012 alone, $10.8 billion was paid out to corn farmers due to losses from
extreme weather, drought, inefficient fertilization, and pests (Barton and Clark,
2014).
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Agricultural researchers have developed a solution to crop loss from pest
species in particular through genetic modification. Specifically, Bt-corn is corn that
has been genetically engineered to express DNA from Bacillus thuringiensis (Bt).
This gram-positive aerobic soil bacterium expresses proteins known as Cry
proteins, which provides the bacteria with its insecticidal characteristics (Sanchis
and Bourguet, 2008). B. thuringiensis is unique in its release of crystalline inclusions
during its sporulation. These crystalline inclusions possess the Cry proteins that are
toxic to insect larvae. Specifically, the crystalline inclusions are dissolved into the
midgut of larval insects releasing Cry proteins. Naturally present stomach enzymes
in the larval midgut synthesize the Cry proteins where it is converted to smaller
polypeptides that are toxic. The toxins now present on the cell membrane of the
larval midgut, forms pores that cause the cells to lyse. Because of this, the larva will
stop feeding and eventually die (Hofte and Whiteley, 1989).
Many strains of B. thuringiensis have been identified and appear to have an
effect on different insects orders. For instance, Cry1Ab and Cry3Bb primarily target
the order Lepidoptera. Specifically, these strains of B. thuringiensis targets the
commonly found European corn borerand corn earworm (Dively and Rose, 2003).
However, toxicity effects have been observed within the species in the orders of
Dipteran and Coleopteran (Hofte and Whiteley, 1989). Despite this, the full toxic
effect of some strains of B. thuringiensis has not yet been identified (Hofte and
Whiteley, 1989).
Bt-corn has become widely adopted by farmers since its introduction nearly
20 years ago, due to its increased production yield and relatively cheap production
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costs (Hofte and Whiteley, 1989). However, with studies observing adverse effects,
concerns have been raised in the scientific community of Bt-corn’s non-target effects
on soil organisms. In particular, non-target effects on species in the order
Hymenoptera have been raised due to their significance in a healthy agricultural
setting. In fact, ants are classified as ecosystem engineers, which means they have a
significant impact on how a habitat is organized. Ants are considered ecosystem
engineers for several reasons with the major one being their capacity to physically
reshape their ecosystem through tunneling. Ants also have high affinity towards
competition and use antibiotic substances to keep invader numbers low and fight
other pest species (Benckiser, 2010). Furthermore, ants are excellent at nutrient
sequestration via their cycling of organic matter with high levels of nitrogen and
carbon, which are crucial for maintaining soil fertility. Lastly, ants are considered
fungal farmers by keeping undesirable fungi at low numbers, and cultivating fungi
that is important for soil fertility and increasing crop yield (Benckiser, 2010).
With the economic and environmental importance of ants in agricultural
ecosystems, I questioned whether differing farming methods have an effect on
species richness and individual abundance. Due to the habitat differences between
polyculture and monoculture farms, I hypothesized that a polyculture establishment
would have higher abundance and species richness than the latter: a farm utilizing
monoculture techniques.
METHODS AND MATERIALS
Study Sites
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Specimens were collected on the 3.6-acre polyculture Lick Run Farm in
Roanoke, Virginia (37.407229, -122.107162), and on the 50-acre monoculture
Michael Beahm Farm in Hollins, Virginia (37.407229, -122.107162). Lick Run Farm
was in an urban environment surrounded by housing developments, and was
serving as a local source of fresh produce for the community. There were two small
plots of cultivation at Lick Run Farm both with leafy vegetables, various peppers,
tomatoes, and sweet corn.
Two rows with approximately 15 corn stalks per row comprised the corn
transect at the organic Lick Run Farm. The first plot (“Fld 1”) was closer to the main
road, whereas the second plot (“Fld 3”) was placed farther away from the main road
(Figure 1). A small dirt pathway led to the second plot, which is where we collected
our specimens. A small field also surrounded this cultivation site (Figure 1). Michael
Beahm farm was in a rural environment with little property development around
cultivation sites (Figure 2). There were two large plots of sweet Bt-corn, which is
where we collected our specimens.
Trapping and Collection
Ants were collected using 50-milliliter screw-cap centrifuge pitfall traps.
Traps were installed on September 19, 2014 and left open for specimen collection
for 12 days. Traps were collected from both farms on October 1, 2014. Two liters of
brine solution was prepared using hot water, salt, vinegar, and regular liquid dish
detergent. A brine solution was substituted for the typical propylene glycol solution
because Lick Run Farm uses organic farming methods with no chemicals. Pitfall
traps were placed on each corner of the corn plot for a total of 8 traps. Pitfall traps
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were also placed directly in the center of the corn patch for a total of 7 traps in the
interior of the corn.
Five pitfall traps were placed randomly in vegetables in the garden plot.
Vegetables that were sampled included red and purple peppers, sweet tomatoes,
lettuce, and grapes. A field gradient was also established around the cultivated site
to sample the natural population of suburban ants. Traps were placed every 10
meters that extended to the corner of the cultivated site, then turned sharply left in
an “L” shape.
Ten traps were placed in two corn plots every 10 meters at Michael Beahm
Farm totaling 20 traps installed. Traps ran parallel to the corn. A field gradient was
also installed at Michael Beahm Farm in a weed patch running parallel to the
cultivated cornfields. Six traps were placed 10 meters apart in order to sample
natural population of ant specimens. By request of Michael Beahm, 4 pitfall traps
were also placed in a white greenhouse building randomly that housed mostly
legumes with some leafy vegetables in order to assess which species of ant had
recently become a greenhouse pest species to Michael Beahm.
Specimen Preservation and Identification
After collection, ants were preserved in alcohol and labeled by date and
location. Specimens were taken back to the lab to be sorted (ant from non-ant) and
pinned for identification. Species were identified using Antweb.org and
dichotomous keys (Fisher and Cover 2007; Holldobler and Wilson 1990; Ellison et
al., 2012). Species were also identified by comparison to vouchers previously
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confirmed by Emily Ogilvy and Dr. Valerie S. Banschbach, Department of Biology,
Saint Michaels College.
Statistical Analyses
Species richness, frequency (ants per trap), relative abundance, and species
diversity were calculated for both the organic farm and the farm with transgenic Bt-
corn. A rarefaction simulation analysis was performed to examine the difference in
species richness present in the overall data set for the organic versus transgenic
corn transects (EstimatesS, Colwell 2013). Rarefaction analysis was performed on
an individual basis. Shannon index (H’) was performed to examine the difference in
species diversity present for both farms. Finally, a t-test was calculated to determine
whether the number of individuals present between the monoculture versus
polyculture study sites were significant or not.
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Figure 1. Aerial photograph of Lick Run Farm. Specimen collection for both field
and corn data occurred in Fld 3.
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Figure 2. Aerial soil map of Michael Beahm Farm. Michael Beahm Farm was
located in rural Hollins, Virginia with little development surrounding it.
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RESULTS
In total, there were 2,214 ants collected on both the organic Lick Run Farm in
Roanoke, and Michael Beahm Farm with transgenic corn in Hollins (Table 1). For
Michael Beahm Farm, there were a total of 357 individuals, and 1,299 individuals
for Lick Run Farm. Of this total, 21 ants species from 13 genera were collected
(Table 2). An outlying colony of 1,015 individuals was collected from Lick Run Farm.
The average number of ants was higher for Lick Run Farm (Table 1; t = -2.66; df =
16; p-value = 0.017). The five most abundant species for the organic Lick Run Farm
were Monomorium emarginatum, Monomoroium viride, Myrmecina americana,
Prenolepis imparis, and Temnothorax ambiguus (Table 2). The most abundant
species for Michael Beahm Farm with transgenic Bt-corn were Lasius alienus, Lasius
umbratus, Pachycondyla chinensis, Prenolepis imparis, and Stenamma brevicorne
(Table 2).
The Shannon diversity index (H’) was higher on the organic Lick Run Farm
(H’ = 2.07) when compared to Michael Beahm Farm (H’ = 1.87) with transgenic Bt-
corn (Table 3). Rarefaction simulation analysis indicated that the organic Lick Run
Farm also had greater species richness than Michael Beahm Farm (Figure 3; Figure
4). Pitfall traps were also placed outside of cultivation to assess differences between
species richness and abundance in a field versus cultivated monoculture or
polyculture. The five most abundant species for the organic Lick Run Farm were
Monomorium viride, Pachycondyla chinensis, Prenolepis imparis, Stenamma
brevicorne, and Temnothorax ambiguus. The five most abundant species for Michael
Beahm Farm were Formica difficilis and Lasius alienus with the same relative
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abundance, Lasius umbratus, Monomorium emarginatum, Prenolepis imparis, and
Stenamma brevicorne (Table 4).
Relative abundance values were calculated for data collected from random
dispersal of pitfall traps in vegetables on the organic Lick Run Farm, and in a hoop
house on Michael Beahm Farm. Values indicate that the top five species in
vegetables on Lick Run Farm are Formica difficilis, Monomorium emarginatum,
Prenolepis imparis, Stenamma brevicorne, and Temnothorax ambiguus (Table 5). The
top five species on Michael Beahm Farm are Formica difficilis, Lasius alienus,
Prenolepis imparis, Stenamma brevicorne, and Temnothorax ambiguus (Table 5).
Table 1. Summary table of total number of ants and traps set with average
number of ants per trap. The organic Lick Run Farm had higher total number of
ants, and higher average number of ants per trap than Michael Beahm Farm with
transgenic Bt-corn.
Site # individuals # traps Mean # ants/trap
Lick Run Farm 1299 17 76
M. Beahm Farm 357 20 17
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Figure 3. Observed ant species richness in corn plots on Lick Run Farm and
Michael Beahm Farm on an individual basis. Observed richness of ants between
the two farms were not different from each other as indicated by the error bars.
Figure 4. Estimated ant species richness in corn plots on Lick Run Farm and
Michael Beahm Farm on an individual basis. Estimated richness of ants between
the two farms indicates Lick Run Farm has higher species richness than Michael
Beahm Farm.
0
2
4
6
8
10
12
14
16
18
0 50 100 150 200 250 300 350 400
ObservedNumberofSpecies
Number of individuals
Lick Run Farm
Beahm Farm
0
5
10
15
20
25
30
0 100 200 300 400 500
EstimatedNumberofSpecies(Chao1)
Number of individuals
Lick Run Farm
Beahm Farm
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Table 2. Ant species relative abundance in the organic (Lick Run Farm) versus
transgenic (Michael Beahm Farm) corn transects. Highlighted cells refer to the
top five most abundant species for each transect.
Ant species Lick Run
Farm
Michael Beahm
Farm
Relative Abundance
Aphaenogaster rudis 0.001 0.015
Camponotus castaneaus 0.001 0.000
Formica difficilis 0.004 0.006
Lasius alienus 0.000 0.122
Lasius umbratus 0.014 0.099
Monomorium
emarginatum
0.055 0.012
Monomorium viride 0.021 0.009
Myrmecina americana 0.781 0.000
Myrmica incompleta 0.000 0.000
Myrmica latifrons 0.000 0.003
Pachycondyla chinensis 0.008 0.163
Pheidole dentate 0.001 0.000
Pheidole flavens 0.001 0.006
Pheidole fulva 0.002 0.038
Pheidole pilifera 0.002 0.012
Prenolepis imparis 0.047 0.353
Solenopsis molesta 0.001 0.003
Stenamma brevicorne 0.013 0.085
Stenamma impar 0.010 0.038
Tapinoma sessile 0.000 0.003
Temnothorax ambiguus 0.038 0.000
Table 3. Shannon diversity index (H’) in the corn plots for organic (Lick Run
Farm) versus transgenic (Michael Beahm Farm). Shannnon diversity index is
higher for the organic Lick Run Farm.
Lick Run Farm Michael Beahm Farm
Shannon index 2.07 1.87
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Table 4. Ant species relative abundance in field transects at the polyculture
Lick Run Farm versus the Michael Beahm Farm with monoculture farming
methods. Highlighted cells refer to the top five most abundant species for each site.
Ant species
Lick Run
Farm
Michael
Beahm Farm
Relative Abundance
Aphaenogaster rudis 0.009 0.053
Formica difficilis 0.074 0.066
Lasius alienus 0.009 0.066
Lasius umbratus 0.009 0.145
Monomorium emarginatum 0.009 0.211
Monomorium viride 0.093 0.000
Myrmica incompleta 0.000 0.026
Myrmica latifrons 0.000 0.013
Myrmica punctiventris 0.000 0.013
Pachycondyla chinensis 0.148 0.000
Pheidole flavens 0.000 0.026
Pheidole pilifera 0.009 0.000
Prenolepis imparis 0.296 0.197
Stenamma brevicorne 0.130 0.026
Stenamma impar 0.000 0.158
Temnothorax ambiguus 0.213 0.000
Table 5. Relative abundance of species in pitfall traps placed in vegetables at
Lick Run Farm and a greenhouse at Michael Beahm Farm. Highlighted cells refer
to the top five most abundant ant species for each farm.
Ant species Lick Run
Veggie
Beahm Hoop
House
Relative Abundance
Aphaenogaster rudis 0.006 0.017
Camponotus castaneus 0.006 0.000
Formica difficilis 0.072 0.157
Lasius alienus 0.011 0.281
Lasius umbratus 0.022 0.058
Monomorium
emarginatum
0.567 0.000
Monomorium viride 0.000 0.008
Pachycondyla chinensis 0.011 0.008
Pheidole flavens 0.044 0.000
Prenolepis imparis 0.067 0.091
Solenopsis molesta 0.006 0.025
Stenamma brevicorne 0.117 0.248
Temnothorax ambiguus 0.072 0.107
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DISCUSSION
This study suggests that diversity and species richness is higher for the
organic, polyculture Lick Run Farm, and there is a statistically significant difference
in abundance of ants in corn plots between Lick Run Farm and Michael Beahm Farm.
On a broader ecological level, this significant difference could be attributed to the
different farming methods of Lick Run Farm and Michael Beahm Farm rather than
the presence of Bt-corn alone. Lick Run farm is an organic polyculture focused
agricultural practice. On the other hand, Michael Beahm Farms’ focus is productivity
via monoculture and utilization of transgenic Bt-corn to efficiently fight pests. A
study conducted in 2013 found statistically significant diversity and richness
differences between a farm utilizing polyculture and a farm utilizing monoculture
(Moreira and Mooney, 2013). They hypothesized this difference could be attributed
to the higher genetic diversity present in polyculture, which attracts aphids and
therefore more resources for ants to utilize (Moreira and Mooney, 2013).
The diversity and richness differences between the two farms could also be
attributed to the dominance of a single invasive ant called the Asian Needle Ant
(Pachycondyla chinensis), which was the second most abundant species at the
Michael Beahm Farm, but was rare at the organic Lick Run Farm (Table 2). It would
be expected to see an abundance of this ant on the organic Lick Run Farm because
an urban environment surrounds it. This is expected because invasive species are
often passengers of disturbance; taking advantage of disturbance to establish as
stated in the driver-passenger hypothesis (MacDougall and Turkington, 2005).
Again, this simple invasive species establishment could be attributed to the
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polyculture versus monoculture. Agricultural researchers are finding that increasing
plant diversity, like polyculture typically does, enhances habitat and resources for
invertebrates, thus creating a stronger ecosystem with a greater variety of
ecological niches making it harder for invasive species to establish (Landis et al.,
2005). Therefore, it is easier for an invasive species like the Asian Needle Ant to
establish because of the low plant diversity and ecological disturbance that are
propagated by monocultures (Landis et al., 2005).
A quick and simple explanation for the abundance and diversity discrepancy
between the two farms would be the presence of transgenic Bt-corn on Michael
Beahm Farm. However, there have been no significant findings that Bt-corn has any
non-target negative affects on ant species (Dively and Rose, 2003). Additionally, a
study comparing the affects of Bt-corn plots without insecticide and non-Bt plots
with insecticide was conducted in 2005. Bt-corn reduces the use of insecticides by
75-100%, and their study concluded that ant and carabid species were the least
affected by insecticide spraying; however, predator species such as spiders and
beetles were decreased by upwards of 75% (Dively and Rose, 2003). This is
concerning ecologically because predator species lower prevalence of insect pests
like the European corn borer and corn earworm in a healthy system, which are the
target species of Bt-corn (Altieri, 1999). Therefore, Bt-corn utilization could have an
indirect positive effect on ants and other predator species because insecticides are
no longer necessary.
For field samples there was some overlap between species found in corn
cultivation and those found in the field. Species that were found in the field but not
19
in the corn for Lick Run Farm were Pachycondyla chinensis, Stenamma brevicorne,
and Monomorium viride. Species that were found in the field but not in the corn for
Michael Beahm Farm were Formica difficilis, Monomorium emarginatum, and
Stenamma impar. It is no surprise to have found the Asian Needle Ant (Pachycondyla
chinensis) on Lick Run Farm since it is an invasive ant that thrives on the
disturbance that an urban environment provides (MacDougall and Turkington,
2005). Due to a strong ecosystem perpetuated by polyculture, the invasive Asian
Needle Ant has likely had difficulty establishing in the cultivated site at Lick Run
Farm (Landis et al., 2005). In addition, the presence of the Asian Needle Ant could be
a driver of species that are present or not. For instance Stenamma brevicorne and
Monomorium viride are potentially good competitors with the Asian Needle Ant
allowing them to sustain their population more than those found in the corn, but not
in the field (Dunn and Guenard, 2010).
This could also be the case for Michael Beahm Farm. Monoculture has
allowed sites of corn cultivation to be more favorable for the invasive Asian Needle
Ant; therefore, ants less tolerant migrate away from the invasive to increase
resources and survival (Dunn and Guenard, 2010). There could also be an
environmental factor that is a contributor to some species being more prevalent in a
natural field versus a cultivated monoculture or polyculture site. For instance, some
species prefer certain types of soil, ground cover, and detritus that may be present
in the field that is not present in the cornfields (Wesson and Wesson, 1940).
In vegetable data for Lick Run Farm and hoop house data for Michael Beahm
Farm, there was some overlap between species that were most abundant.
20
Interestingly, However, ant species that were not found to be abundant in the corn
plots were found to be abundant in the hoop house and vegetable plots. These
species include Formica difficilis and Stenamma brevicorne for Lick Run Farm
vegetable plots, and Formica difficilis and Temnothorax ambiguus for Michael Beahm
Farm hoop house (Table 3). On Michael Beahm Farm, this could be attributed to the
decline in Pachycondyla chinensis abundance and the possible intolerance of
Formica difficilis and Temnothorax ambiguus to the Asian Needle Ant (Dunn and
Guenard, 2010). The dominance of Temnothorax ambiguus and Formica difficilis in
vegetable plots rather than corn plots at Lick Run Farm could be attributed to the
type of soil the vegetables were growing in. For instance, Formica difficilis prefer dry
soil with vegetable debris, and Stenamma brevicorne prefer moist soil (Wesson and
Wesson, 1940). Therefore, they would be more abundant in vegetables with the
type of soil preferred by the organism.
In conclusion, my hypothesis was accepted with the finding of significant
statistical differences in the number of individuals between the polyculture Lick Run
Farm and monoculture Michael Beahm Farm (t = -2.66; df = 16; p-value = 0.017).
This could be attributed to the differing farming methods used by the farmers.
Overall, transgenic Bacillus thuringiensus (Bt) corn does not appear to have adverse
affects on non-target organisms in the Hymenoptera family based on literature
(Dively and Rose, 2003). The foundation has been established for further research
on Lick Run Farm and Michael Beahm Farm with the identification of species that
are present. Therefore, more research can increase knowledge in a time of
21
agricultural shifting in the United States of the impacts of certain agricultural
practices on important bioindicator species like ants.
ACKNOWLEDGEMENTS
The Roanoke College Environmental Studies Program including Dr. Valerie S.
Banschbach and Dr. Katherine P. O’Neill, Lick Run Farm in Roanoke, Virginia, and
Michael Beahm Farm in Hollins, Virginia. Dr. Matthew Petersen in the Roanoke
College Biology Department. Dr. Valerie S. Banschbach and the Roanoke College
Environmental Studies Program provided supplies and equipment necessary for
this project.
LITERATURE CITED
Altieri, Miguel A. 1999 Ecological role of biodiversity in agroecosystems. Agriculture,
Ecosystems, and the Environment. 74: 19-31.
Barton, Brooke and Sarah Elizabeth Clark. 2014 Water and climate risks facing U.S.
corn production: how companies and investors can cultivate sustainability.
Ceres. 1-71.
Benckiser, Gero. 2010 Ants and sustainable agriculture: A review. Agron. Sustain.
Dev. 30: 191-199.
Colwell, Robert K. 2013. EstimateS: Statistical estimation of species richness and
shared species from samples. Version 9 and earlier. User’s Guide and
application. http://purl.oclc.org/estimates.
Crawford, Jim. 2011 Lick Run Farm 1st Work Day. Vimeo Video.
https://vimeo.com/31646855.
22
Daly, Timothy, and G. David Buntin. 2005 transgenic corn for Lepidopteran control
on non-target arthropods. Environ. Entomol. 34: 1292-1301.
Dively, G.P., and R. Rose. 2003 Effects of Bt transgenic and conventional insecticide
control on the non-target natural enemy community in sweet corn.
International Symposium on Biological Control of Arthropods. 265-274.
Dunn, Robert R., and Benoit Guenard. 2010 A new (old), invasive ant in the
hardwood forests of eastern North America and its potentially widespread
impacts. Plos One.
Ellison, Aaron M., Nicholas J. Gotelli, and Elizabeth J. Farnsworth. 2012 A field guide
to the ants of New England. Yale University Press. 1-398.
Fisher, Brian L., and Stefan P. Cover. 2007 Ants of North America: A guide to the
genera. University of California Press. 1-194.
Hofte, Herman and H.R. Whiteley. 1989 Insecticidal crystal proteins of Bacillus
thuringiensis. Microbiological Reviews. 53: 242-255.
Holldobler, Bert, and Edward O. Wilson. 1992 The ants. The Belknap Press of
Harvard University Press. 1-732.
Irving, Sarah, Rob Harrison, and Mary Rayner. 2001 Ethical consumerism –
democracy through the wallet. Journal of Research for Consumers. 3: 1-20.
Kjar, Daniel. 2009 The ant community of a riparian forest in Dyke Marsh Preserve,
Fairfax County, Virginia, and a checklist of Mid-Atlantic Formicinae.
Banisteria. 33: 3-17.
23
Landis, Douglas A., Fabian D. Menalled, and Alejandro C. Costamagna. 2005
Manipulating plant resources to enhance beneficial arthropods in
agricultural landscapes. Weed Science. 53: 902-908.
MacDougall, Andrew S., and Roy Turkington. 2005 Are invasive species the drivers
or passengers of change in degraded ecosystems? Ecology. 86: 42-55.
Moreira, Xaoquin and Kailen A. Mooney. 2013 Influence of plant genetic diversity on
interactions between higher trophic levels. Biology Letters. 9: 130-133.
Postma, Johannes A., Jonathan P. Lynch. 2012 Complementarity in root architecture
for nutrient uptake in ancient maize/ bean and maize/bean/squach
polycultures. Ann. Bot. 2: 521-534.
Sanchis, Vincent and Denis Bourguet. 2008 Bacillus thuringiensis: Applications in
agriculture and insect resistance management. A review. Agron. Sustain. Dev.
28: 11-20.
Vandermeer, John H. 2011 The ecology of agroecosystems Jones and Bartlett
Publishers, LLC.
Wesson, Laurence G., and Robert G. Wesson. 1940 A collection of ants from
southcentral Ohio. American Midland Naturalist. 24: 89-103.
Zhang, Wei, Taylor H. Ricketts, Claire Kremen, Karen Carney, and Scott M. Swinton.
2007 Ecosystem services and dis-services to agriculture. Ecological
Economics. 8: 1-8.

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FINAL THESIS PAPER

  • 1. 1 A Study Comparing Ant Richness and Diversity Between Two Agricultural Ecosystems: Polyculture versus Monoculture Abagail E. Davis and Valerie S. Banschbach (Advisor) Department of Environmental Studies, Roanoke College Salem, Virginia 29 April 2015
  • 2. 2 ABSTRACT In today’s agriculture, there are two types of farming methods: monoculture and polyculture. Monoculture is born out of the industrial system and is when one crop is planted on a single plot of land (Vandermeer, 2011). In contrast, polyculture is typically smaller operations, organic, and the practice of planting a variety of crops on a single plot of land (Vandermeer, 2011). Both agricultural methods produce differing ecosystems. Typically found on large monoculture operations are transgenic organisms like Bt-Corn. Corn is a crucial cash crop for small-scale and large, industrial farmers with the United States leading the way in production and exportation (Barton and Clark, 2014). However, with global climate change constantly looming, farmers have observed an increase in corn pest species like the European corn borer and corn earworm (Dively and Rose, 2003). Therefore, agricultural researchers have developed a genetically engineered corn crop composed of Bacillus thuringiensus, a gram-positive soil bacterium that has insecticidal characteristics to combat pest issues on large-scale monocultures (Sanchis and Bourguet, 2008). This study explored the question of whether differing agricultural practices of monoculture and polyculture affect species richness and diversity of ants on two farms: an polyculture farm called Lick Run Farm, and a monoculture farm with transgenic corn called Michael Beahm Farm. Results indicate species richness and diversity is higher on Lick Run Farm than on Michael Beahm Farm. Higher diversity and richness is likely based on the monoculture At Michael Beahm Farm versus polyculture agricultural methods at Lick Run Farm.
  • 3. 3 INTRODUCTION In an age of agricultural revitalization in the United States, consumers are beginning to put more thought into their food purchases, and gravitating more towards food items that have been cultivated sustainably (Irving et al., 2001). The current agricultural paradigm in the United States is fueled by the industrial system with goals of growth, profit, and productivity. This paradigm is driven by monoculture, or the cultivation of a single crop on a plot of land (Vandermeer, 2011). Monoculture is favored in the industrial agricultural ecosystem because it is less work than its alternative, and it speaks to productivity (Vaandermeer, 2011). An alternative to monoculture; on the other hand, is polyculture. Polyculture is the cultivation of many varieties of crops on a single plot of land together at the same time (Vandermeer, 2011). Monoculture and polyculture, in essence, are differing habitats. The biology and ecology for both forms of agriculture are differing. This difference will be illustrated using the two farm sites used for this study: Lick Run Farm and Michael Beahm Farm. Lick Run Farm is an organic, but not certified, 3-acre polyculture farm in the heart of the city of Roanoke, Virginia. It is surrounded by urban environment with houses in every direction. Prior to becoming Lick Run Farm, the property was home to a nursery called Crowell Nursery (Crawford, 2011). The soil prior to Lick Run Farm ownership was degraded and lacking important nutrients. In an effort to restore the soil, Rick Williams, the owner of Lick Run Farm, began planting cover crops. Organic, polyculture farmers often use cover crops, or crops planted to add
  • 4. 4 nutrients back to the soil and protect against erosion. Examples of cover crops are legumes, various grasses, and grains (Zhang et al., 2007). Williams also adds mulch and compost to his soil for extra enrichment. Williams utilizes plant fragrances for pest management and to attract important pollinators as well. Another important regime Williams uses on Lick Run Farm is the three sisters method. The three sisters method utilizes intercropping of maize, beans, and squash for efficient nutrient sequestration and uptake by the soil and the plant, while also deterring pests. For instance, what nitrogen is taken up by the corn the beans replenish back into the soil (Postma and Lynch, 2012). Michael Beahm Farm is a 200-230 acre, monoculture farm in rural Hollins, Virginia with 50-acres of cropland growing transgenic sweet Bt-corn. The Bt-corn was not sprayed with any herbicides or pesticides. Michael Beahm Farm has been in the Beahm family for several years now, and has always been operating as a larger- scale monoculture operation. Corn is a crucial agricultural commodity to the United States comprising nearly 94 million acres of farmland in 2013. It is the most economically valuable crop estimating a total production value of $67 billion (Barton and Clark, 2014). Additionally, the United States is the number one producer and exporter of corn comprising 36% of the global output. Unfortunately, natural events out of humankind’s control are posing serious threats to the success of corn in the United States. In 2012 alone, $10.8 billion was paid out to corn farmers due to losses from extreme weather, drought, inefficient fertilization, and pests (Barton and Clark, 2014).
  • 5. 5 Agricultural researchers have developed a solution to crop loss from pest species in particular through genetic modification. Specifically, Bt-corn is corn that has been genetically engineered to express DNA from Bacillus thuringiensis (Bt). This gram-positive aerobic soil bacterium expresses proteins known as Cry proteins, which provides the bacteria with its insecticidal characteristics (Sanchis and Bourguet, 2008). B. thuringiensis is unique in its release of crystalline inclusions during its sporulation. These crystalline inclusions possess the Cry proteins that are toxic to insect larvae. Specifically, the crystalline inclusions are dissolved into the midgut of larval insects releasing Cry proteins. Naturally present stomach enzymes in the larval midgut synthesize the Cry proteins where it is converted to smaller polypeptides that are toxic. The toxins now present on the cell membrane of the larval midgut, forms pores that cause the cells to lyse. Because of this, the larva will stop feeding and eventually die (Hofte and Whiteley, 1989). Many strains of B. thuringiensis have been identified and appear to have an effect on different insects orders. For instance, Cry1Ab and Cry3Bb primarily target the order Lepidoptera. Specifically, these strains of B. thuringiensis targets the commonly found European corn borerand corn earworm (Dively and Rose, 2003). However, toxicity effects have been observed within the species in the orders of Dipteran and Coleopteran (Hofte and Whiteley, 1989). Despite this, the full toxic effect of some strains of B. thuringiensis has not yet been identified (Hofte and Whiteley, 1989). Bt-corn has become widely adopted by farmers since its introduction nearly 20 years ago, due to its increased production yield and relatively cheap production
  • 6. 6 costs (Hofte and Whiteley, 1989). However, with studies observing adverse effects, concerns have been raised in the scientific community of Bt-corn’s non-target effects on soil organisms. In particular, non-target effects on species in the order Hymenoptera have been raised due to their significance in a healthy agricultural setting. In fact, ants are classified as ecosystem engineers, which means they have a significant impact on how a habitat is organized. Ants are considered ecosystem engineers for several reasons with the major one being their capacity to physically reshape their ecosystem through tunneling. Ants also have high affinity towards competition and use antibiotic substances to keep invader numbers low and fight other pest species (Benckiser, 2010). Furthermore, ants are excellent at nutrient sequestration via their cycling of organic matter with high levels of nitrogen and carbon, which are crucial for maintaining soil fertility. Lastly, ants are considered fungal farmers by keeping undesirable fungi at low numbers, and cultivating fungi that is important for soil fertility and increasing crop yield (Benckiser, 2010). With the economic and environmental importance of ants in agricultural ecosystems, I questioned whether differing farming methods have an effect on species richness and individual abundance. Due to the habitat differences between polyculture and monoculture farms, I hypothesized that a polyculture establishment would have higher abundance and species richness than the latter: a farm utilizing monoculture techniques. METHODS AND MATERIALS Study Sites
  • 7. 7 Specimens were collected on the 3.6-acre polyculture Lick Run Farm in Roanoke, Virginia (37.407229, -122.107162), and on the 50-acre monoculture Michael Beahm Farm in Hollins, Virginia (37.407229, -122.107162). Lick Run Farm was in an urban environment surrounded by housing developments, and was serving as a local source of fresh produce for the community. There were two small plots of cultivation at Lick Run Farm both with leafy vegetables, various peppers, tomatoes, and sweet corn. Two rows with approximately 15 corn stalks per row comprised the corn transect at the organic Lick Run Farm. The first plot (“Fld 1”) was closer to the main road, whereas the second plot (“Fld 3”) was placed farther away from the main road (Figure 1). A small dirt pathway led to the second plot, which is where we collected our specimens. A small field also surrounded this cultivation site (Figure 1). Michael Beahm farm was in a rural environment with little property development around cultivation sites (Figure 2). There were two large plots of sweet Bt-corn, which is where we collected our specimens. Trapping and Collection Ants were collected using 50-milliliter screw-cap centrifuge pitfall traps. Traps were installed on September 19, 2014 and left open for specimen collection for 12 days. Traps were collected from both farms on October 1, 2014. Two liters of brine solution was prepared using hot water, salt, vinegar, and regular liquid dish detergent. A brine solution was substituted for the typical propylene glycol solution because Lick Run Farm uses organic farming methods with no chemicals. Pitfall traps were placed on each corner of the corn plot for a total of 8 traps. Pitfall traps
  • 8. 8 were also placed directly in the center of the corn patch for a total of 7 traps in the interior of the corn. Five pitfall traps were placed randomly in vegetables in the garden plot. Vegetables that were sampled included red and purple peppers, sweet tomatoes, lettuce, and grapes. A field gradient was also established around the cultivated site to sample the natural population of suburban ants. Traps were placed every 10 meters that extended to the corner of the cultivated site, then turned sharply left in an “L” shape. Ten traps were placed in two corn plots every 10 meters at Michael Beahm Farm totaling 20 traps installed. Traps ran parallel to the corn. A field gradient was also installed at Michael Beahm Farm in a weed patch running parallel to the cultivated cornfields. Six traps were placed 10 meters apart in order to sample natural population of ant specimens. By request of Michael Beahm, 4 pitfall traps were also placed in a white greenhouse building randomly that housed mostly legumes with some leafy vegetables in order to assess which species of ant had recently become a greenhouse pest species to Michael Beahm. Specimen Preservation and Identification After collection, ants were preserved in alcohol and labeled by date and location. Specimens were taken back to the lab to be sorted (ant from non-ant) and pinned for identification. Species were identified using Antweb.org and dichotomous keys (Fisher and Cover 2007; Holldobler and Wilson 1990; Ellison et al., 2012). Species were also identified by comparison to vouchers previously
  • 9. 9 confirmed by Emily Ogilvy and Dr. Valerie S. Banschbach, Department of Biology, Saint Michaels College. Statistical Analyses Species richness, frequency (ants per trap), relative abundance, and species diversity were calculated for both the organic farm and the farm with transgenic Bt- corn. A rarefaction simulation analysis was performed to examine the difference in species richness present in the overall data set for the organic versus transgenic corn transects (EstimatesS, Colwell 2013). Rarefaction analysis was performed on an individual basis. Shannon index (H’) was performed to examine the difference in species diversity present for both farms. Finally, a t-test was calculated to determine whether the number of individuals present between the monoculture versus polyculture study sites were significant or not.
  • 10. 10 Figure 1. Aerial photograph of Lick Run Farm. Specimen collection for both field and corn data occurred in Fld 3.
  • 11. 11 Figure 2. Aerial soil map of Michael Beahm Farm. Michael Beahm Farm was located in rural Hollins, Virginia with little development surrounding it.
  • 12. 12 RESULTS In total, there were 2,214 ants collected on both the organic Lick Run Farm in Roanoke, and Michael Beahm Farm with transgenic corn in Hollins (Table 1). For Michael Beahm Farm, there were a total of 357 individuals, and 1,299 individuals for Lick Run Farm. Of this total, 21 ants species from 13 genera were collected (Table 2). An outlying colony of 1,015 individuals was collected from Lick Run Farm. The average number of ants was higher for Lick Run Farm (Table 1; t = -2.66; df = 16; p-value = 0.017). The five most abundant species for the organic Lick Run Farm were Monomorium emarginatum, Monomoroium viride, Myrmecina americana, Prenolepis imparis, and Temnothorax ambiguus (Table 2). The most abundant species for Michael Beahm Farm with transgenic Bt-corn were Lasius alienus, Lasius umbratus, Pachycondyla chinensis, Prenolepis imparis, and Stenamma brevicorne (Table 2). The Shannon diversity index (H’) was higher on the organic Lick Run Farm (H’ = 2.07) when compared to Michael Beahm Farm (H’ = 1.87) with transgenic Bt- corn (Table 3). Rarefaction simulation analysis indicated that the organic Lick Run Farm also had greater species richness than Michael Beahm Farm (Figure 3; Figure 4). Pitfall traps were also placed outside of cultivation to assess differences between species richness and abundance in a field versus cultivated monoculture or polyculture. The five most abundant species for the organic Lick Run Farm were Monomorium viride, Pachycondyla chinensis, Prenolepis imparis, Stenamma brevicorne, and Temnothorax ambiguus. The five most abundant species for Michael Beahm Farm were Formica difficilis and Lasius alienus with the same relative
  • 13. 13 abundance, Lasius umbratus, Monomorium emarginatum, Prenolepis imparis, and Stenamma brevicorne (Table 4). Relative abundance values were calculated for data collected from random dispersal of pitfall traps in vegetables on the organic Lick Run Farm, and in a hoop house on Michael Beahm Farm. Values indicate that the top five species in vegetables on Lick Run Farm are Formica difficilis, Monomorium emarginatum, Prenolepis imparis, Stenamma brevicorne, and Temnothorax ambiguus (Table 5). The top five species on Michael Beahm Farm are Formica difficilis, Lasius alienus, Prenolepis imparis, Stenamma brevicorne, and Temnothorax ambiguus (Table 5). Table 1. Summary table of total number of ants and traps set with average number of ants per trap. The organic Lick Run Farm had higher total number of ants, and higher average number of ants per trap than Michael Beahm Farm with transgenic Bt-corn. Site # individuals # traps Mean # ants/trap Lick Run Farm 1299 17 76 M. Beahm Farm 357 20 17
  • 14. 14 Figure 3. Observed ant species richness in corn plots on Lick Run Farm and Michael Beahm Farm on an individual basis. Observed richness of ants between the two farms were not different from each other as indicated by the error bars. Figure 4. Estimated ant species richness in corn plots on Lick Run Farm and Michael Beahm Farm on an individual basis. Estimated richness of ants between the two farms indicates Lick Run Farm has higher species richness than Michael Beahm Farm. 0 2 4 6 8 10 12 14 16 18 0 50 100 150 200 250 300 350 400 ObservedNumberofSpecies Number of individuals Lick Run Farm Beahm Farm 0 5 10 15 20 25 30 0 100 200 300 400 500 EstimatedNumberofSpecies(Chao1) Number of individuals Lick Run Farm Beahm Farm
  • 15. 15 Table 2. Ant species relative abundance in the organic (Lick Run Farm) versus transgenic (Michael Beahm Farm) corn transects. Highlighted cells refer to the top five most abundant species for each transect. Ant species Lick Run Farm Michael Beahm Farm Relative Abundance Aphaenogaster rudis 0.001 0.015 Camponotus castaneaus 0.001 0.000 Formica difficilis 0.004 0.006 Lasius alienus 0.000 0.122 Lasius umbratus 0.014 0.099 Monomorium emarginatum 0.055 0.012 Monomorium viride 0.021 0.009 Myrmecina americana 0.781 0.000 Myrmica incompleta 0.000 0.000 Myrmica latifrons 0.000 0.003 Pachycondyla chinensis 0.008 0.163 Pheidole dentate 0.001 0.000 Pheidole flavens 0.001 0.006 Pheidole fulva 0.002 0.038 Pheidole pilifera 0.002 0.012 Prenolepis imparis 0.047 0.353 Solenopsis molesta 0.001 0.003 Stenamma brevicorne 0.013 0.085 Stenamma impar 0.010 0.038 Tapinoma sessile 0.000 0.003 Temnothorax ambiguus 0.038 0.000 Table 3. Shannon diversity index (H’) in the corn plots for organic (Lick Run Farm) versus transgenic (Michael Beahm Farm). Shannnon diversity index is higher for the organic Lick Run Farm. Lick Run Farm Michael Beahm Farm Shannon index 2.07 1.87
  • 16. 16 Table 4. Ant species relative abundance in field transects at the polyculture Lick Run Farm versus the Michael Beahm Farm with monoculture farming methods. Highlighted cells refer to the top five most abundant species for each site. Ant species Lick Run Farm Michael Beahm Farm Relative Abundance Aphaenogaster rudis 0.009 0.053 Formica difficilis 0.074 0.066 Lasius alienus 0.009 0.066 Lasius umbratus 0.009 0.145 Monomorium emarginatum 0.009 0.211 Monomorium viride 0.093 0.000 Myrmica incompleta 0.000 0.026 Myrmica latifrons 0.000 0.013 Myrmica punctiventris 0.000 0.013 Pachycondyla chinensis 0.148 0.000 Pheidole flavens 0.000 0.026 Pheidole pilifera 0.009 0.000 Prenolepis imparis 0.296 0.197 Stenamma brevicorne 0.130 0.026 Stenamma impar 0.000 0.158 Temnothorax ambiguus 0.213 0.000 Table 5. Relative abundance of species in pitfall traps placed in vegetables at Lick Run Farm and a greenhouse at Michael Beahm Farm. Highlighted cells refer to the top five most abundant ant species for each farm. Ant species Lick Run Veggie Beahm Hoop House Relative Abundance Aphaenogaster rudis 0.006 0.017 Camponotus castaneus 0.006 0.000 Formica difficilis 0.072 0.157 Lasius alienus 0.011 0.281 Lasius umbratus 0.022 0.058 Monomorium emarginatum 0.567 0.000 Monomorium viride 0.000 0.008 Pachycondyla chinensis 0.011 0.008 Pheidole flavens 0.044 0.000 Prenolepis imparis 0.067 0.091 Solenopsis molesta 0.006 0.025 Stenamma brevicorne 0.117 0.248 Temnothorax ambiguus 0.072 0.107
  • 17. 17 DISCUSSION This study suggests that diversity and species richness is higher for the organic, polyculture Lick Run Farm, and there is a statistically significant difference in abundance of ants in corn plots between Lick Run Farm and Michael Beahm Farm. On a broader ecological level, this significant difference could be attributed to the different farming methods of Lick Run Farm and Michael Beahm Farm rather than the presence of Bt-corn alone. Lick Run farm is an organic polyculture focused agricultural practice. On the other hand, Michael Beahm Farms’ focus is productivity via monoculture and utilization of transgenic Bt-corn to efficiently fight pests. A study conducted in 2013 found statistically significant diversity and richness differences between a farm utilizing polyculture and a farm utilizing monoculture (Moreira and Mooney, 2013). They hypothesized this difference could be attributed to the higher genetic diversity present in polyculture, which attracts aphids and therefore more resources for ants to utilize (Moreira and Mooney, 2013). The diversity and richness differences between the two farms could also be attributed to the dominance of a single invasive ant called the Asian Needle Ant (Pachycondyla chinensis), which was the second most abundant species at the Michael Beahm Farm, but was rare at the organic Lick Run Farm (Table 2). It would be expected to see an abundance of this ant on the organic Lick Run Farm because an urban environment surrounds it. This is expected because invasive species are often passengers of disturbance; taking advantage of disturbance to establish as stated in the driver-passenger hypothesis (MacDougall and Turkington, 2005). Again, this simple invasive species establishment could be attributed to the
  • 18. 18 polyculture versus monoculture. Agricultural researchers are finding that increasing plant diversity, like polyculture typically does, enhances habitat and resources for invertebrates, thus creating a stronger ecosystem with a greater variety of ecological niches making it harder for invasive species to establish (Landis et al., 2005). Therefore, it is easier for an invasive species like the Asian Needle Ant to establish because of the low plant diversity and ecological disturbance that are propagated by monocultures (Landis et al., 2005). A quick and simple explanation for the abundance and diversity discrepancy between the two farms would be the presence of transgenic Bt-corn on Michael Beahm Farm. However, there have been no significant findings that Bt-corn has any non-target negative affects on ant species (Dively and Rose, 2003). Additionally, a study comparing the affects of Bt-corn plots without insecticide and non-Bt plots with insecticide was conducted in 2005. Bt-corn reduces the use of insecticides by 75-100%, and their study concluded that ant and carabid species were the least affected by insecticide spraying; however, predator species such as spiders and beetles were decreased by upwards of 75% (Dively and Rose, 2003). This is concerning ecologically because predator species lower prevalence of insect pests like the European corn borer and corn earworm in a healthy system, which are the target species of Bt-corn (Altieri, 1999). Therefore, Bt-corn utilization could have an indirect positive effect on ants and other predator species because insecticides are no longer necessary. For field samples there was some overlap between species found in corn cultivation and those found in the field. Species that were found in the field but not
  • 19. 19 in the corn for Lick Run Farm were Pachycondyla chinensis, Stenamma brevicorne, and Monomorium viride. Species that were found in the field but not in the corn for Michael Beahm Farm were Formica difficilis, Monomorium emarginatum, and Stenamma impar. It is no surprise to have found the Asian Needle Ant (Pachycondyla chinensis) on Lick Run Farm since it is an invasive ant that thrives on the disturbance that an urban environment provides (MacDougall and Turkington, 2005). Due to a strong ecosystem perpetuated by polyculture, the invasive Asian Needle Ant has likely had difficulty establishing in the cultivated site at Lick Run Farm (Landis et al., 2005). In addition, the presence of the Asian Needle Ant could be a driver of species that are present or not. For instance Stenamma brevicorne and Monomorium viride are potentially good competitors with the Asian Needle Ant allowing them to sustain their population more than those found in the corn, but not in the field (Dunn and Guenard, 2010). This could also be the case for Michael Beahm Farm. Monoculture has allowed sites of corn cultivation to be more favorable for the invasive Asian Needle Ant; therefore, ants less tolerant migrate away from the invasive to increase resources and survival (Dunn and Guenard, 2010). There could also be an environmental factor that is a contributor to some species being more prevalent in a natural field versus a cultivated monoculture or polyculture site. For instance, some species prefer certain types of soil, ground cover, and detritus that may be present in the field that is not present in the cornfields (Wesson and Wesson, 1940). In vegetable data for Lick Run Farm and hoop house data for Michael Beahm Farm, there was some overlap between species that were most abundant.
  • 20. 20 Interestingly, However, ant species that were not found to be abundant in the corn plots were found to be abundant in the hoop house and vegetable plots. These species include Formica difficilis and Stenamma brevicorne for Lick Run Farm vegetable plots, and Formica difficilis and Temnothorax ambiguus for Michael Beahm Farm hoop house (Table 3). On Michael Beahm Farm, this could be attributed to the decline in Pachycondyla chinensis abundance and the possible intolerance of Formica difficilis and Temnothorax ambiguus to the Asian Needle Ant (Dunn and Guenard, 2010). The dominance of Temnothorax ambiguus and Formica difficilis in vegetable plots rather than corn plots at Lick Run Farm could be attributed to the type of soil the vegetables were growing in. For instance, Formica difficilis prefer dry soil with vegetable debris, and Stenamma brevicorne prefer moist soil (Wesson and Wesson, 1940). Therefore, they would be more abundant in vegetables with the type of soil preferred by the organism. In conclusion, my hypothesis was accepted with the finding of significant statistical differences in the number of individuals between the polyculture Lick Run Farm and monoculture Michael Beahm Farm (t = -2.66; df = 16; p-value = 0.017). This could be attributed to the differing farming methods used by the farmers. Overall, transgenic Bacillus thuringiensus (Bt) corn does not appear to have adverse affects on non-target organisms in the Hymenoptera family based on literature (Dively and Rose, 2003). The foundation has been established for further research on Lick Run Farm and Michael Beahm Farm with the identification of species that are present. Therefore, more research can increase knowledge in a time of
  • 21. 21 agricultural shifting in the United States of the impacts of certain agricultural practices on important bioindicator species like ants. ACKNOWLEDGEMENTS The Roanoke College Environmental Studies Program including Dr. Valerie S. Banschbach and Dr. Katherine P. O’Neill, Lick Run Farm in Roanoke, Virginia, and Michael Beahm Farm in Hollins, Virginia. Dr. Matthew Petersen in the Roanoke College Biology Department. Dr. Valerie S. Banschbach and the Roanoke College Environmental Studies Program provided supplies and equipment necessary for this project. LITERATURE CITED Altieri, Miguel A. 1999 Ecological role of biodiversity in agroecosystems. Agriculture, Ecosystems, and the Environment. 74: 19-31. Barton, Brooke and Sarah Elizabeth Clark. 2014 Water and climate risks facing U.S. corn production: how companies and investors can cultivate sustainability. Ceres. 1-71. Benckiser, Gero. 2010 Ants and sustainable agriculture: A review. Agron. Sustain. Dev. 30: 191-199. Colwell, Robert K. 2013. EstimateS: Statistical estimation of species richness and shared species from samples. Version 9 and earlier. User’s Guide and application. http://purl.oclc.org/estimates. Crawford, Jim. 2011 Lick Run Farm 1st Work Day. Vimeo Video. https://vimeo.com/31646855.
  • 22. 22 Daly, Timothy, and G. David Buntin. 2005 transgenic corn for Lepidopteran control on non-target arthropods. Environ. Entomol. 34: 1292-1301. Dively, G.P., and R. Rose. 2003 Effects of Bt transgenic and conventional insecticide control on the non-target natural enemy community in sweet corn. International Symposium on Biological Control of Arthropods. 265-274. Dunn, Robert R., and Benoit Guenard. 2010 A new (old), invasive ant in the hardwood forests of eastern North America and its potentially widespread impacts. Plos One. Ellison, Aaron M., Nicholas J. Gotelli, and Elizabeth J. Farnsworth. 2012 A field guide to the ants of New England. Yale University Press. 1-398. Fisher, Brian L., and Stefan P. Cover. 2007 Ants of North America: A guide to the genera. University of California Press. 1-194. Hofte, Herman and H.R. Whiteley. 1989 Insecticidal crystal proteins of Bacillus thuringiensis. Microbiological Reviews. 53: 242-255. Holldobler, Bert, and Edward O. Wilson. 1992 The ants. The Belknap Press of Harvard University Press. 1-732. Irving, Sarah, Rob Harrison, and Mary Rayner. 2001 Ethical consumerism – democracy through the wallet. Journal of Research for Consumers. 3: 1-20. Kjar, Daniel. 2009 The ant community of a riparian forest in Dyke Marsh Preserve, Fairfax County, Virginia, and a checklist of Mid-Atlantic Formicinae. Banisteria. 33: 3-17.
  • 23. 23 Landis, Douglas A., Fabian D. Menalled, and Alejandro C. Costamagna. 2005 Manipulating plant resources to enhance beneficial arthropods in agricultural landscapes. Weed Science. 53: 902-908. MacDougall, Andrew S., and Roy Turkington. 2005 Are invasive species the drivers or passengers of change in degraded ecosystems? Ecology. 86: 42-55. Moreira, Xaoquin and Kailen A. Mooney. 2013 Influence of plant genetic diversity on interactions between higher trophic levels. Biology Letters. 9: 130-133. Postma, Johannes A., Jonathan P. Lynch. 2012 Complementarity in root architecture for nutrient uptake in ancient maize/ bean and maize/bean/squach polycultures. Ann. Bot. 2: 521-534. Sanchis, Vincent and Denis Bourguet. 2008 Bacillus thuringiensis: Applications in agriculture and insect resistance management. A review. Agron. Sustain. Dev. 28: 11-20. Vandermeer, John H. 2011 The ecology of agroecosystems Jones and Bartlett Publishers, LLC. Wesson, Laurence G., and Robert G. Wesson. 1940 A collection of ants from southcentral Ohio. American Midland Naturalist. 24: 89-103. Zhang, Wei, Taylor H. Ricketts, Claire Kremen, Karen Carney, and Scott M. Swinton. 2007 Ecosystem services and dis-services to agriculture. Ecological Economics. 8: 1-8.