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Goldenrod Galls

The impact of gall-forming parasites on plant growth and the study of viability of parasitic

offspring

Alicia N. Cleaver

Ecology and Evolution 007

Professor Tepe

22 November 2017
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Abstract

Parasites are infamous in the world of biology for their negative effects on host

organisms. The average person may think of any number of microscopic worms or other

creatures when parasites are mentioned. However, parasites of all types can infect plant species.

This study aims to analyze one particular relationship between gall-forming insects in the

common goldenrod plant. Eurosta solidaginis and Rhopalomyia solidaginis are flies which

oviposit their larva into the flesh of the goldenrod. The larva form galls, or hollow pockets, that

sustain them until their adult emergence. This relationship poses a few questions including if

density of goldenrods impacts gall presence, if galls affect plant growth, and if the width of the

gall is related to the fate of the gall maker. Data analysis has revealed that all three relationships

hold significance.

Introduction

Parasitism is a non-mutual relationship in which one species benefits at the expense of

another. It occurs in nearly every community on earth. Parasites gain nutrients from their hosts

and may even benefit by receiving shelter and a habitat to grow and reproduce in. A unique

parasitic interaction occurs between gall-making insects and Solidago Canadensis, commonly

known as the goldenrod plant. This plant-insect relationship is of interest because it exemplifies

the direct impact that parasites have on their hosts and vice versa.

Eurosta solidaginis is one of the insects analyzed in this observational study. It is a fly

that induces ball-shaped galls on the stem of the goldenrod plant (Abrahamson et al, 1989;

Figure 1A). It does this by secreting chemicals that induce rapid cell division and enlargement of

plant tissue (Weis & Berenbaum 1989). The larva of Eurosta feed on the inner tissue of the plant

and remain in the chamber for about 50 weeks until emergence (Yahnke 2006). The other fly
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species observed in this study, Rhopalomyia solidaginis, induces galls and has a life cycle that is

much the same as Eurosta. However, Rhopalomyia oviposit on near the tip of growing plants and

cause galls to form in the apical region (Weis & Berenbaum 1989; Figure 2A).

Figure 1A. A gall on the stem of a golden- Figure 2A. A gall on the apical region

rod. Photo by Thomas Wilson. of a goldenrod. Photo by Richard Orr.

Both species of parasitic flies benefit from the shelter and nutrients provided by the

goldenrod plants. However, they are not entirely immune to other parasites and predators.

Natural enemies of Eurosta and Rhopalomyia include parasitoid wasps and avian predators such

as the black-capped chickadee and downy woodpecker (Abrahamson et al., 1989). In general, the

wasps oviposit into smaller galls because their walls are thinner; the ovipositor must be injected

through the wall and onto the living body of the gall-maker larva. In contrast, it is believed that

birds crack into larger galls because they are more visible. This may be true also because smaller

galls are more likely to contain parasitic wasp larva which is less nutritious than gall-making

insect larva (Yahnke 2006).

Survival from wasp parasitism and bird predation of Eurosta and Rhopalomyia ultimately

depends on gall size. Smaller galls are susceptible to parasitoid oviposition, but larger galls are
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more attractive to birds. Therefore, gall-making insects in medium-sized galls have the highest

rates of survival; there seems to be a force of stabilizing selection acting on the size of galls

(Abrahamson et al., 1989). Based off this information, it is hypothesized that gall width is related

to fate of the gall maker.

Previous research has indicated that gall presence from Eurosta and Rhopalomyia

impacts plant growth. Both stem and apical galls negatively affected goldenrod plants in a study

performed in Pennsylvania. Apical and stem gall-bearing plants showed a significant decrease in

height when compared to non-gall-bearing plants. There were also significant reductions in seed

reproductive allocation (Hartnett & Abrahamson 1979). It is hypothesized that gall presence does

impact plant growth in this study.

Methods

Data collection occurred on the 1st of November, 2017 at the UC Center for Field Studies.

The center is a multi-acre mix of woodlands, creeks, wetlands, and prairie. The temperature was

about 45º Fahrenheit. The prairie area of the reserve served as the main site for collection. It is

adjacent to a small creek and directly next to a grass path. Several groups established a 10m long

transect in the field, perpendicular to the path. A tape measure was laid upon the transect and

anchored down. Starting at the one meter mark, 1m x 1m quadrats were established on each side

of the tape using another smaller tape measure. A total of 10 1m x 1m quadrats were established

in each group.

The groups then counted the number of parasitized and unparasitized goldenrods in each

quadrat. The number of apical galls and stem galls was counted and recorded. A random

assortment of five of each apical parasitized, stem parasitized, and unparasitized goldenrods were

selected and measured. This was done by starting at the ground and extending to the highest
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point on the plant with the small tape measure. Finally, any dead goldenrods with galls from the

previous season were located and collected. They were examined and placed into different

categories. Unsuccessful galls were those which did not appear to have any openings from larval

emergence. Successful galls were those which had a small, drill bit-size opening signifying larval

emergence. Wasp parasitized and bird parasitized galls were also placed into separate categories

based on appearance of the opening.

The data was then compiled into an excel document and analyzed using ANOVA single

factor tests and the descriptive statistics analysis function. The ANOVA single factor test is a

statistical method used to compare the null hypothesis against the alternative; the null states that

the means of each sample are equal, whereas the alternative states they are not the same. The

density and growth of unparasitized, apical gall, and stem gall goldenrods was tested in an

ANOVA. A T-test was generated for the density of stem and apical galls to determine whether

the means of the two groups were statistically significant.

The success of gall makers and their fate was also analyzed using the ANOVA method.

The descriptive statistics function was then generated for each of the aforementioned categories

in an effort to find the means and standard errors. This mean numbers were placed into bar

graphs to show the relationship between each of the groups.

Results

Table 1. ANOVA test for total density of unparasitized, apical and stem gall-bearing goldenrods.

ANOVA
Source of
Variation SS df MS F P-value F crit
9.25782E-
Between Groups 256471.423 2 128235.711 276.739351 95 3.00554033
Within Groups 424920.224 917 463.380833

Total 681391.647 919

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45
40
35
30
Density (m2)

25
20
15
10
5
0
1 2 3
Unparasitized Stem Apical

Figure 1. Bar graph of mean density comparisons of unparasitized, apical and stem gall-bearing

goldenrods.

Table 1A. T-test of density between stem galls and apical galls.

t-Test: Two-Sample Assuming Unequal Variances

Variable 1 Variable 2
Mean 0.237458194 4.026578073
Variance 0.396444524 25.39929125
Observations 299 301
Hypothesized Mean
Difference 0
df 309
-
t Stat 12.94271273
P(T<=t) one-tail 3.28076E-31
t Critical one-tail 1.649799826
P(T<=t) two-tail 6.56151E-31
t Critical two-tail 1.967670885
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The ANOVA depicted in Table 1 is a test for total density of unparasitized, stem and gall

bearing goldenrods. It generated the values (F= 276.74, df= 2, p= 9.26E-95). The bar graph in

Figure 1 illustrates the difference in mean density between the three variables. Unparasitized

goldenrods were most prevalent in the regions sampled with a mean density of about 37 plants

per meter squared. Apical galls were the next most dense with roughly 4 galls per meter squared.

Finally, stem galls had a density of about 0.2 galls per meter squared. Error bars from the graph

also indicate statistical significance from lack of overlap. The T-test in Table 1A reiterates that

there is a level of significance between the density of apical and stem galls.

Table 2. ANOVA test for height of unparasitized, apical and stem gall-bearing goldenrods.

ANOVA
Source of
Variation SS df MS F P-value F crit
Between Groups 328047.0301 2 164023.515 70.56177598 0 2.999471607
Within Groups 5583539.78 2402 2324.537794

Total 5911586.811 2404

140

120

100
Height (m)

80

60

40

20

0
1 2 3
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Figure 2. Bar graph comparing the mean heights of unparasitized, apical and stem gall-bearing

goldenrods.

The ANOVA depicted in Table 2 compared the heights of unparasitized, apical

parasitized and stem parasitized goldenrods. It generated the values (F= 70.56, df= 2, p= 0). The

bar graph in Figure 2 illustrates the differences in mean height between the three variables.

Unparasitized goldenrods had a mean height of 121.55cm. Goldenrods with galls on their stems

had a mean height of 106.23cm, and those with apical galls had a mean height of 97.04cm. Error

bars from the graph also indicate statistical significance from lack of overlap.

Table 3. ANOVA test for successful and unsuccessful insect emergence in relation to wasp

parasitism and bird predation. Gall width was the final factor examined.

Source of
Variation SS df MS F P-value F crit
9.12614E-
Between Groups 97.3030358 3 32.434345 12.189956 07 2.705838051
Within Groups 239.4668961 90 2.66074329

Total 336.7699319 93
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6
Gall Width (mm)

0
1 2 3 4
Unsuccesful Successful Wasp Parasitism Bird Predation

Figure 3. Bar graph of mean successful and unsuccessful gall emergence in relation to

disturbance from wasp parasitism and bird predation. Gall width was the final factor examined.

The ANOVA depicted in Table 3 analyzed several factors that influence fate of gall

makers. Unsuccessful and successful larval emergence was determined in relation to gall width.

The gall widths of plants that were parasitized by wasps and fed on by birds were also measured.

The ANOVA generated the values (F= 12.19, df= 3, p= 9.13E-07). These mean results from the

data are depicted in bar graph form in Figure 3. Unsuccessful galls had a mean width of 3.3mm

and successful had a mean width of 4.20mm. Wasp parasitized and bird predated had mean

widths of 6.51mm and 5.29 mm, respectively. Error bars from the bar graph also indicate

statistical significance from lack of overlap.

Discussion

The data revealed several interesting points about the questions posed. The extremely low

p-value of 9.26E-95 from the ANOVA test in Table 1 indicates that a significant difference does

exist between the density of unparasitized, stem and gall goldenrods. It can be assumed that galls

will be present in areas with high goldenrod populations. Figure 1 also shows that apical galls are
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far more common than stem galls. It is possible that some unknown force decreased the

population of Eurosta in the area tested and therefore decreased the amount of stem galls. More

observation in future years could be performed to determine if patterns of stem and apical gall-

bearing plants fluctuate.

The data analysis for heights of unparasitized, apical, and stem parasitized plants

generated notable results, as well. The ANOVA in Table 2 generated a p-value of 0 which means

that the null hypothesis can be rejected; there is a significant difference in heights between the

factors tested. This may be attributed to the fact that goldenrods are directly impacted by the

presence of apical and stem gall-bearing goldenrods. Goldenrods with galls generally do not

grow as high as their afflicted counterparts (Hartnett & Abrahamson 1979). Figure 2 also

confirms the fact that galls impact plant growth. An interesting further study could examine why

galls impact goldenrod growth so markedly. Perhaps apical galls stunt growth more than stem

galls because they restrict the formation of tall upper leaves that aid in photosynthesis.

Data showed that gall width is related to fate maker. The ANOVA from Table 3 had a p-

value of 9.13E-07 meaning statistical significance does exist between the factors tested.

Interestingly enough, the data seems to contradict previous studies that state that wasps generally

oviposit into smaller galls because their walls are thinner (Abrahamson et al, 1989). The parasitic

wasps laid their larva in galls with a mean width of 6.51mm, which is larger than the vast

majority of gall measurements taken in this study. The mean width for bird predation was

slightly lower than that of the wasp. In general, it did seem that larger galls had greater success

rates (Figure 3). This supports the hypothesis that gall width is related to fate of the gall maker.

Further studies could be designed to examine the minimum and maximum gall widths that
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successfully support larva. Very small galls may not be able to provide the nourishment and

shelter needed, whereas very large galls would make emergence difficult.

Works Cited

Abrahamson, Warren G., et al. “Variation in selection pressures on the goldenrod gall fly and the

competitive interactions of its natural enemies.” Oecologia, vol. 79, no. 1, 1989, pp. 15–

22.

Hartnett, David C., and Warren G. Abrahamson. “The Effects of Stem Gall Insects on Life

History Patterns in Solidago Canadensis.” Ecology, vol. 60, no. 5, 1979, pp. 910–917.

Orr, Richard. A Goldenrod Bunch Gall Midge gall in Howard Co., Maryland.2014, photograph,

Maryland Biodiversity Project.

Weis AE & Berenbaum MR. 1989. Herbivorous insects and green plants. In: Plant-animal

interactions (Abrahamson WG, ed.), McGraw-Hill Book Co., New York.

Wilson, Thomas. A Goldenrod Gall Fly Gall in Baltimore City, Maryland. 2013, photograph,

Maryland Biodiversity Project.

Yahnke, C. J. “Testing Optimal Foraging Theory: Using Bird Predation on Goldenrod

Galls.” The American Biology Teacher, vol. 68, no. 8, 0 Oct. 2006, pp. 471–475.