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The effects of selection of goldenrod galls’ height and diameter on Eurosta solidaginis

Abstract

We examined the effects of selective agents acting on the height and diameter of Eurosta

solidaginins goldenrod galls. We compared the goldenrod galls that contained live

Eurosta with the galls of dead Eurosta, examining height and diameter. We hypothesized

that selection favors lower galls that are less visible to avian predators and larger galls

that are less likely to be successfully penetrated by the wasp Eurytoma gigantean due to

their larger size. We observed significantly larger diameters of live Eurosta’s galls than

dead Eurosta’s galls but did not find significant differences between the heights of galls

of live and dead Eurosta.

Introduction

A measure of fitness in the goldenrod gall fly, Eurosta solidaginis, is larval

survivorship. Gall size has been implicated in differential survivorship of Eurosta larva

(Weis and Abrahamson, 1985), which makes it a likely candidate to experience selective

pressures. We examined the relationship between gall diameter and larval survivorship,

and gall height and larval survivorship to gain insight into the selective effects of

parasitism and predation on Eurosta galls.

Eurosta oviposits in June, inserting an egg into the terminal bud of the goldenrod

Solidago altissima (Weis and Abrahamson, 1986). The larva tunnels into the stem

immediately below the apical meristem (Weis and Abrahamson, 1985) where it releases

chemicals that stimulate the host goldenrod to form a gall in which the larva will live

until emergence the next May (Abrahamson and Heinrich, web source: Bucknell

University). The gall forms from plant tissue, but the larva’s genotype affects gall size
(Weis and Abrahamson, 1986). The heritability of gall size in Eurosta supports the theory

that Eurosta have been selected to stimulate gall formation and that the gall response is a

manipulation of the goldenrod host plant by the larva (Weis and Abrahamson, 1986).

Predators and parasites account for the majority of larval mortality (Weis and

Abrahamson, 1985, 1986). Eurosta’s natural enemies, birds and parasitic wasps,

differentially affect larva survival based on goldenrod phenotype (size measured by

diameter) and exert both directional and stabilizing selection (Weis and Abrahamson,

1986). Directional selection increases the proportion of a certain phenotype in a

population as a result of greater fitness or reproductive success of individuals with the

phenotype. Stabilizing selection selects towards the mean of a phenotype that varies on a

continuum when displaying phenotypic extremes causes decreased fitness. Stabilizing

selection narrows the variance of a certain phenotype in a population. Weis and

Abrahamson (1985) have shown that small gall size increases the likelihood of successful

attack by the Eurytoma gigantea wasp, causing a selective pressure toward larger gall

size. The selective pressures at work are not unidirectional; large size increases

susceptibility to attack by the Downy Woodpecker and the Black-capped chickadee,

producing an opposing selective pressure toward small gall size (Weis and Abrahamson,

1986). In our examination of selective pressures acting on Eurosta, we expected to see

directional selection favoring larger gall diameter and to see stabilizing selection

modifying the effects of directional selection by selecting against very large and very

small gall diameter.

Gall height is influenced by the height of the goldenrod at the time of oviposition

and the goldenrod’s rate of growth. Eurosta females have the capacity to discriminate
when choosing goldenrods for oviposition based on signals that indicate host quality i.e.

characteristics of the plant that increase her larvae’s chance of survival (Anderson et al.

1989). Horner and Abrahamson (1999) found that goldenrod genotype significantly

affects gall height at the time of oviposition. Anderson et al. (1989) observed that Eurosta

females select goldenrods that were growing significantly faster and were taller than the

rest of the population at the time of oviposition, though 3 and 4 weeks later they were

growing slower than the rest of the population (the heights remained significantly

different). They identified a probable cause for the selection of faster growing

goldenrods: faster growing goldenrods are more responsive to the stimulus of the Eurosta

larva. Anderson et al. proposed that the subsequent reduction in growth rate was caused

by allocation of limited resources to gall formation. Anderson et al. (1989) also found

greater larval mortality in goldenrod genotypes that are generally more resistant to gall

formation; therefore, Eurosta females do not only select goldenrods that are more

susceptible to gall formation but also select goldenrods that increase the chances of larvae

surviving. If heritable traits of Eurosta females affect gall height and gall height affects

larval survival, natural selection can act on gall height. We predicted that natural selection

favors shorter galls.

We examined the effect of goldenrod gall height and gall diameter on

survivorship of Eurosta in order to identify if selective pressures were acting on Eurosta,

influencing gall phenotype. We measured gall height and diameter and categorized galls

as containing alive or dead Eurosta larva. We analyzed the differences in gall phenotype

between the alive and dead samples for height and diameter to determine if there were

significant differences between the two samples that might indicate selection.
We explored the effects of selection on gall diameter by comparing the difference

between the diameter of galls containing live larvae and the diameter of galls containing

dead larvae. We predicted that galls containing live Eurosta larvae would have greater

diameter than galls in which the Eurosta larvae had died because small galls are more

vulnerable to parasite attack (Weis and Abrahamson, 1985). Galls have finished growth

during the period of E. gigantea oviposition (Weis and Abrahamson, 1985). The smaller

diameter of galls containing dead larvae does not reflect terminated growth after larval

death, but would be consistent with directional selection against small gall size. We also

predicted stabilizing selection as a result of the opposing selective pressures exerted by

avian predators that are more likely to attack large diameter galls (Weis and Abrahamson,

1985). We predicted that natural selection acts on gall height. We hypothesized that it

favors decreased gall height as a result of increased bird predation if taller galls are more

attractive or apparent to birds.

Materials and Methods

Lab instructors collected galls on January 9th through January 13th 2006 at Hildacy

Farm. We collected data on January 19, 2006: we measured gall height in centimeters

with a measuring tape held straight from the base of the stem where the golden rod was

cut at ground level to the midpoint of the gall. We measured gall diameter in millimeters

at the widest part of the gall with digital calipers which were first zeroed in closed

position. The pith of the galls were scored vertically with a knife and opened without

damaging live larva inside.

We examined the contents of galls and categorized the gall as containing dead

Eurosta solidaginis if a living Eurosta larva was absent. We tried to ascertain fungal or
bacterial cause of death under the microscope. We categorized Eurosta as alive if we

found and identified a living Eurosta larva. If the gall contained an inhabitant other than a

Eurosta larva we categorized the Eurosta as dead. When possible, we identified the

organism present as one of the parasitic species that invade Eurosta galls such as wasps

Eurytoma gigantea and Eurytoma obtusiventris or Mordellistena. We recorded physical

characteristics of the gall and its contents when pertinent. For example, after categorizing

a gall as containing dead Eurosta, we noted that the gall was cracked and that a bird

possibly caused the death. Data about a gall from a goldenrod gall moth was discarded as

not pertinent to our experiment on goldenrod Eurosta galls. We included 173 galls in our

data.

We used the Shapiro-Wilk W test to test for normality in height by larval status

and diameter by larval status. We used a test for non-parametric comparison, the

Wilcoxon rank-sum test, to compare the median height of galls containing living Eurosta

with the median height of galls containing dead Eurosta because the data for heights of

galls containing live Eurosta was not normal (the data for galls containing dead Eurosta

was normal). Similarly, we used the Wilcoxon test to compare the median diameter of

galls containing live Eurosta with that of galls containing dead Eurosta because the data

for diameters of galls containing live Eurosta was not normal (data for galls containing

dead Eurosta was normal). The Wilcoxon test was an appropriate test to ascertain

whether the differences between the median height and diameter of live larvae and the

median height and diameter of dead larvae were significantly different. The Wilcoxon test

ascertains the probability that the results of experimentation are due to random variation

seen from sample size.


Results

We examined the diameter and height of 173 goldenrod galls induced by Eurosta

Solidaginis. The galls were divided into two samples which were comprised of 120 galls

containing live Eurosta and 53 galls of dead Eurosta. Galls containing live Eurosta had

greater diameter than galls of dead Eurosta. Gall height was similar across larval status.

We observed galls containing live Eurosta (median=21.7 mm, upper IQR=1.8 mm, lower

IQR=2.47 mm; n=120) had significantly greater median diameter than galls of dead

Eurosta (median=19.1 mm, upper IQR=1.6 mm, lower IQR=2.3 mm; n=53, P <.0001;

Figure 1). The slightly greater median height of galls of dead Eurosta (median=82.6 cm,

upper IQR=17.0 cm, lower IQR=14.85 cm; n=53) was not significantly different from the

median height of galls containing live Eurosta (median=80 cm, upper IQR=11.75 cm,

lower IQR=8.15 cm; n=120, P=.474; Figure 2).

Discussion

Weis and Abrahamson (1986) showed that the mean size of selected individuals’

galls, those that survived and presumably reproduced, was significantly greater than the

mean of the whole gene pool. E. gigantea prey on larvae after the galls have reached

their full size, therefore small gall size is not a result of larva death terminating growth

(Weis and Abrahamson, 1986). Their data indicated that the effect of upward pressure on

gall size by E. gigantea outweighed that of the opposing pressure toward smaller gall size

that birds exert. Their data also supported the presence of stabilizing selection acting to

reduce variance. Similarly our data showed significantly greater median gall size for live

Eurosta galls than for galls of dead Eurosta, confirming selection toward larger gall

diameter. The distribution of diameters of live Eurosta galls appears to reflect stabilizing
selection but we did not perform addition statistical tests to identify the characteristics of

the non-normal distributions.

A dearth of literature on the effects of selection on Eurosta gall height, especially

when compared to the preponderance of research on the effects of selection on gall size,

suggests that selection does not affect gall height as strongly as it affects gall diameter.

The absence of selection on gall height might result from an unfulfilled requirement for

selection—that in order for natural selection to occur there must be genetically based

variation in the trait (Purrington et al. Swarthmore College Bio 2 lecture).

Gall height must be a heritable character of Eurosta for selection to act on it.

Selection could act on height if females’ selection of goldenrods for oviposition, which is

likely a heritable trait, affected gall height and if height influenced the survivorship of

larvae. Horner and Abrahamson (1999) investigated goldenrod response to simulated

drought stress and found that goldenrod genotype modulates and causes varying degrees

of response. In some genotypes, goldenrods flowered significantly earlier under control

conditions than under drought stress but others displayed a reverse trend, while some

experienced no significant effects. Ovipositing females discerned between drought-

simulation and control plants, selecting more control plants. Females also responded

significantly to genotype in their selection, but there was no significant effect of the

interaction between treatment and genotype, despite the demonstrated effect of genotype

on reaction to treatment.

We found no significant difference between the median height of galls of dead

Eurosta and the median height of live Eurosta galls. We propose that because the signals

used by Eurosta in goldenrod selection do not incorporate the effects of the interaction of
genetics and the environment that selective pressures do not have a strong effect on gall

height. Height must be heritable for selection to act and, though the ability to select more

suitable goldenrods is likely a heritable trait, goldenrod genotypes interact with the

environment in diverse ways, which most likely reduces the effect of selective pressures

(if any) on goldenrod height.

Literature Cited

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Teaching and Research.
http://www.facstaff.bucknell.edu/abrahmsn/solidago/main.html>. [accessed 2005
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