Am. Midl. Nat.

163:173–185

Pattern and Consequences of Floral Herbivory in Four

Sympatric Ipomoea Species
DEXTER R. SOWELL1 AND LORNE M. WOLFE
Department of Biology, Georgia Southern University, P.O. Box 8042, Statesboro 30460

ABSTRACT.—Flowers are expected to be under strong selection pressure to be attractive to

pollinators. Yet, floral traits that serve in pollinator attraction may inadvertently attract
animals that consume flowers. We expected animals to utilize floral cues in targeting plants to
attack so for three field seasons we examined florivory in four sympatric Ipomoea species that
differed in their pollination syndrome (flower color and shape) as well as their geographic
origin (native vs. introduced). Most of the damage to Ipomoea flowers was caused by larval
Spodoptera spp. that consumed all or parts of the corolla, pistil and stamens. There was no
overall difference in damage levels to flowers of the bee or hummingbird-pollinated plants.
However, florivores exhibited a strong preference for the native, bumble bee-pollinated I.
cordatotriloba relative to the remaining three non-native Ipomoea species. The non-native,
hummingbird-pollinated I. hederifolia and I. quamoclit consistently experienced the lowest
florivory rates. There were reproductive consequences to florivory as damaged flowers had
reduced fruit set. Seed set was not influenced by corolla damage, but was reduced when
flowers suffered damage to reproductive organs. The results of our study suggest that flower-
eating insects are not focusing in on traits associated with different pollination syndromes,
but are preferentially visiting the native Ipomoea species, and this may contribute to the
successful invasion of introduced Ipomoea plants that suffer reduced damage levels.

INTRODUCTION
Flowers of animal-pollinated plants should be under strong selection to be attractive to
pollinators owing to their central role in reproduction (Waser, 1978; Johnson and Dafni,
1998; Wolfe and Krstolic, 1999; Kawagoe and Suzuki, 2004). Features such as flower color,
size, number and type of reward all participate in attracting pollinators for pollen
dissemination and receipt (Waser and Price, 1981; Schemske and Horovitz, 1984; Nilsson,
1988; Harder, 1990; Herrera, 1993; Hodges, 1995; Johnson et al., 1995; Campbell et al., 1996;
Conner and Rush, 1996; Galen, 1996). Angiosperms that utilize the same pollinator or
pollinator guild (e.g., hummingbirds) often have flowers that have similar suites of floral
traits referred to as pollination syndromes (Faegri and van der Pijl, 1979; Fenster et al.,
2004). Yet, traits that serve to differentially attract pollinators may at the same time
differentially attract organisms that feed on flowers (Armbruster, 1997; Matter et al., 1999;
Galen and Cuba, 2001; Leege and Wolfe, 2002; Canela and Sazima, 2003; Held and Potter,
2004). Flowers, therefore, act as resources for both mutualistic pollinators and antagonistic
predators (Armstrong and Marsh, 1997; Wolfe, 1997; Irwin and Brody, 1998; Krupnick and
Weis, 1999; Galen and Cuba, 2001; Irwin et al., 2004).
Florivores can have obvious direct effects on plant fitness if they damage the reproductive
structures (stamens and pistils), resulting in the loss of gametes (Breedlove and Ehrlich,
1968; Cunningham, 1995; Krupnick and Weis, 1999). Damage to attractive structures
(corolla and calyx) can also indirectly affect reproductive success if these damaged flowers
receive fewer pollinator visits, or visits of lower quality (Irwin and Brody, 1999; Krupnick et

1
Corresponding author present address: Department of Biology, University of Virginia, P.O. Box
400328, Charlottesville 22904; email: dsowell@virginia.edu; Telephone: (434) 982-5218; FAX: (434) 982-
5628

173
174 THE AMERICAN MIDLAND NATURALIST 163(1)

al., 1999; Mothershead and Marquis, 2000; McCall, 2006; Pohl et al., 2006). Despite the
impact on plant reproductive success, little is known about how florivores respond to
variation in floral traits that clearly function in pollinator attraction (Leege and Wolfe, 2002;
Irwin et al., 2004).
The goal of this study was to evaluate whether sympatric Ipomoea (morning glory) species
with similar pollination syndromes also experience similar florivory intensities (proportion of
flowers damaged), and what role native vs. introduced status plays in shaping florivory intensity.
In the southeast US, we observed lepidopteran larvae (predominately Spodoptera spp.) causing
damage to flowers, but not harming foliage, of four common Ipomoea species. These vining
plants produce flowers that differ in appearance and can be grouped into two different
pollination syndromes (Faegri and van der Pijl, 1979). Ipomoea hederifolia and I. quamoclit have
red-orange and red flowers, respectively, with narrow tubular corollas, dilute nectar and are
predominately visited by hummingbirds and sulfur butterflies (hereafter referred to as
‘hummingbird-pollinated’ Ipomoea). Ipomoea hederacea and I. cordatotriloba have blue and purple
flowers, respectively, with larger flared corolla openings, concentrated nectar and are
predominately visited by bumble bees (hereafter referred to as ‘bumble bee-pollinated’
Ipomoea) (Wolfe and Sowell, 2006). In addition to differences in floral morphology, the four
Ipomoea species differ in their origins. Whereas Ipomoea cordatotriloba is native to the southeast
US, the remaining three species were introduced from tropical America (Godfrey and Wooten,
1981) and are considered noxious weeds by the United States Department of Agriculture
(2001). A popular idea in the invasion biology literature is that invasiveness results when
introduced species escape their enemies (e.g., Elton, 1958; Crawley, 1987; Wolfe, 2002). Thus,
these four species provide an excellent system to evaluate not only the possible antagonistic
effects of attraction and attack, but whether species are native or introduced, because
individuals of the four Ipomoea species grow intermingled at several local sites.
Specifically, we addressed the following questions: (1) Does the intensity and pattern of
florivory differ between the Ipomoea species? If so, does pollination syndrome or origin
(native/introduced) explain variation in florivory among the focal Ipomoea species? Are
these patterns consistent between years? (2) What flower parts, attractive (corolla) or
reproductive structures (stamens, pistil), are damaged or consumed by florivores? (3) What
is the relative effect of damage to attractive vs. reproductive structures on fruit and seed set?

METHODS
Study system.—This study was conducted on four Ipomoea species common in southeast
Georgia (Bulloch County) in 1999, 2000 and 2005: I. cordatotriloba Dennst., I. hederifolia L., I.
quamoclit L. and I. hederacea (L.) Jacq., with nomenclature following Austin and Huáman
(1996), with subgeneric delineation following Miller et al. (1999). A recent molecular study
(Miller et al., 1999) upholds previous morphologically-defined systematic placement of the
bumble bee-pollinated I. hederacea and I. cordatotriloba into different subgenera, Ipomoea and
Eriospermum, respectively. The hummingbird-pollinated I. hederifolia and I. quamoclit, are
closely related and belong to subgenus Quamoclit.
Ipomoea cordatotriloba is a native perennial, whereas the remaining three species are non-
native annuals (Godfrey and Wooten, 1981). In southeast Georgia, the species flower from
Jul. to first frost (typically early Nov.) and exhibit strongly congruent seasonal flowering
phenologies (Wolfe and Sowell, 2006). Individual flowers remain open for less than one day,
with anthesis beginning at or shortly before daybreak (Wolfe and Sowell, 2006). Fruits
mature in 3–4 wk, and the dehiscent fruit capsules contain a maximum of six seeds in I.
hederacea, and four seeds in I. hederifolia, I. quamoclit and I. cordatotriloba.
2010 SOWELL & WOLFE: HERBIVORY IN IPOMOEA 175

Measuring the magnitude of florivory.—Our goal was to determine the magnitude and
consequences of florivory for each of the four Ipomoea species. We assessed florivory intensity
(proportion of flowers with damage) at approximately weekly intervals throughout each
flowering season at seven frequently disturbed roadsides. All sites were separated by at least
several miles, and were alongside roads or railways. Each site contained up to several
hundred flowers of each Ipomoea species daily. The specific sites used varied among years
owing to availability of flowering material because some sites were mown or received
herbicide application from state transportation workers before or shortly after our surveying
began. These sites were eliminated from further surveying within the study year and not
included in analyses. Sites surveyed by year were: 1999-Hunter’s Pointe, Hwy 24, Hwy 67;
2000-Hunter’s Pointe, Hopeulikit, Hwy 80E; 2005-Mill Creek, Railroad. Each year surveying
started in early Aug. and continued until late Oct. or early Nov., encompassing peak
flowering and the majority of the seasonal flowering period of the four Ipomoea species
(Wolfe and Sowell, 2006).
Although a slightly different sampling method was used each year, the overall approach
provided a robust measure of florivory intensity. In 1999 we performed exhaustive censuses
of all intact and damaged Ipomoea flowers throughout the entire site (no subsampling within
sites) on each survey date to obtain site-level florivory intensity. In 1999 and 2005, we
distinguished between three types of floral damage: (1) attractive structures (corolla only),
(2) reproductive structures (pistil and stamens only) and (3) whole flower (attractive and
reproductive structures) to determine the patterns of floral damage among Ipomoea species.
In 2000 and 2005, we counted the number of intact and damaged flowers within
permanent plots distributed regularly throughout the sites. In 2000, 5 m2 plots were
established in early Jul., just before flowering began, but after growing vines were clearly
visible, and were distributed approximately every 10 m linearly throughout the sites. In
2005, 1 m2 plots were established along transects approximately every 2 m throughout the
sites. Within plots, all flowers of each Ipomoea species were counted, and recorded as being
intact or damaged. We summed intact and damaged flowers across plots within a site to yield
an estimate of site-level florivory intensity (proportion of surveyed flowers with damage) for
each Ipomoea species. Over the course of the three study years, we surveyed approximately
48,000 flowers among the four Ipomoea species (Table 1).
For all years, each census was performed at midday, after florivore activity ceased (Sowell
and Wolfe, pers. obs.), to provide a realistic assessment of florivory intensity on a daily
cohort of flowers.
Measuring the consequences of florivory.—We determined the consequences of florivory by
comparing fruit and seed production in damaged and undamaged flowers. Our protocol
was to locate and tag naturally damaged flowers along with adjacent undamaged flowers
(within 30 cm) around midday after florivore activity had ceased. Small paper tags placed on
the pedicel identified flowers as damaged or undamaged. We returned 3–4 wk later to
collect the mature fruit. Flowers that did not set fruit retained tags on their pedicels and
were recorded as unsuccessful flowers.
We used three sites in 1999 (Hunter’s Pointe, Hwy 24, Hwy 67) and one site in 2005 (Mill
Creek) to quantify fruit and seed set. Fruit set for Ipomoea cordatotriloba was excluded from
analysis in 1999 because too few flowers were produced on days when we tagged flowers to
follow through fruit set, most likely due to drought conditions. In 1999, we did not
differentiate type of floral damage (attractive, reproductive organs) in our damage
treatment, tagging 109 damaged and 109 undamaged flowers across five days in early Sept.
and early Oct. In 2005, we tagged 105 damaged flowers and 347 undamaged flowers across
176 THE AMERICAN MIDLAND NATURALIST 163(1)

TABLE 1.—Number of flowers surveyed across Ipomoea species and study years

Species 1999 2000 2005 Species Total

I. hederifolia 17,718 2375 805 20,898

I. quamoclit 12,439 862 1362 14,663
I. hederacea 2493 233 142 2868
I. cordatotriloba 6143 2769 648 9560
Year total 38,793 6239 2957 47,989

14 d from early Sept. through early Nov. and recorded the type of floral damage (attractive,
reproductive structures) to determine the fate of flowers that had their reproductive organs
damaged relative to those that received damage only to attractive structures.
Statistical analyses.—All analyses were conducted with JMP version 3.2.1 (SAS, 1997).
Nonparametric analyses were used when assumptions of normality and heteroscedasticity
could not be met by data transformation. We utilized ranks of florivory intensity (proportion
of flowers damaged) as our response variable, in a Scheirer-Ray-Hare extension of Kruskal-
Wallis test (Sokal and Rohlf, 1995).
For analyses on florivory intensity, we included the following in the model: species, site,
species by site interaction and number of flowers available to florivores of each Ipomoea species
on each census day as a covariate (Table 1). Florivory intensity did not vary by sampling date
within a season, and we have excluded sampling date from the analyses. Separate analyses,
utilizing the same model stated above, were conducted for each of the 3 y because a different
suite of sites was sampled each year. We tested for differences in rank florivory intensity
between the individual Ipomoea species with post-hoc analyses, examining the affects of origin
and pollination syndrome, with significance values adjusted by a sequential Bonferroni
correction (Rice, 1989). The frequency of damage to different floral tissues (attractive,
reproductive structures) in the four species was analyzed with a chi-square test.
Fruit and seed set in three Ipomoea species in 1999, and four Ipomoea species in 2005, was
analyzed with the effect of species, flower damage (yes, no) and the species—flower damage
interaction as effect variables. For fruit set, we utilized a two-way logistic regression, with fruit
maturation (yes, no) as the response variable. For seed set, we utilized an ordinal logistic
regression, with seed number as the response variable. For both the logistic and ordinal
regression we report likelihood ratio chi-squares (X2). Throughout the manuscript we
present the mean 6 SE.

RESULTS
MAGNITUDE OF FLORIVORY

Florivory was ubiquitous in the three study years. Pooling across the three study years, we
observed florivory intensity (percentage of flowers damaged) that ranged from a low of 0.96
6 0.25% in Ipomoea hederifolia, to a high of 6.44 6 0.91% in I. cordatotriloba (Fig. 1). There
were significant differences in florivory intensity among the Ipomoea species in 1999 and
2000, but not significantly so in 2005, following sequential Bonferroni correction (Table 1).
Post-hoc analyses revealed that levels of florivory did not differ between the two pollination
syndromes. However, the native I. cordatotriloba suffered higher florivory rates compared to
the three non-native focal Ipomoea (Table 2).
The degree of damage we observed was consistent among sites within years, except for
1999, where sites were significantly different for flower damage levels after Bonferroni
2010 SOWELL & WOLFE: HERBIVORY IN IPOMOEA 177

TABLE 2.—The analysis of florivory intensity on four Ipomoea species in southeastern Georgia. Two-way
ANCOVA of ranks (Scheirer-Ray-Hare extension of the Kruskal-Wallis test) testing the effects of species,
site, species by site interaction, and daily number of flowers available within species as a covariate, on the
proportion of flowers damaged daily by florivores. Post-hoc analyses with species pooled by their origin
(native, introduced) and pollination syndrome (hummingbird-, bumble bee-pollinated) follow below
the species entry. For each year, P values significant following a sequential Bonferroni correction
are bolded

Source df SS H P,

1999
Species 3 112,128.30 63.35 ,0.0001
Origin 1 46,965.85 26.53 ,0.0001
Pollination syndrome 1 394.87 0.22 0.64
Site 2 17,326.65 9.79 0.01
Species 3 site 6 45,356.30 25.62 ,0.0002
Flower number 1 2776.51 1.57 0.21
Error 126 52,467.42
2000
Species 3 50,935.17 45.01 ,0.0001
Origin 1 39,612.82 35.01 ,0.0001
Pollination syndrome 1 1684.74 1.49 0.23
Site 2 6227.36 5.50 0.07
Species 3 site 6 6952.86 6.14 0.41
Flower number 1 1523.04 1.35 0.25
Error 100 52,735.04
2005
Species 3 984.40 4.62 0.03
Origin 1 906.81 4.26 0.04
Pollination syndrome 1 216.77 1.02 0.32
Site 1 46.79 0.22 0.64
Species 3 site 3 148.69 0.70 0.87
Flower number 1 555.55 2.61 0.11
Error 38 7606.80

correction (Table 2). Furthermore, the florivory intensity within each Ipomoea species was
not affected by the number of flowers available in each site on a given day (Table 1).
The most frequently attacked floral structure was the corolla (Fig. 2), with over 60% of
damaged flowers receiving damage that varied from patchy removal of corolla tissue to the
loss of the entire corolla (pers. obs.). The pattern of damage to floral parts within each
species was similar between 1999 and 2005. Overall, approximately 50% of damaged flowers
had their reproductive organs damaged. Stamens and pistils, when damaged, were usually
completely consumed by florivores. In both 1999 and 2005, the frequency of damage to
the different floral structures varied among the four Ipomoea species (Fig. 3; 1999: X2 5
17.26, df 5 6, P , 0.008; 2005: X2 5 18.54, df 5 6, P , 0.005).

REPRODUCTIVE CONSEQUENCES OF FLORIVORY

Fruit set.—Overall, flower damage significantly reduced fruit set in the Ipomoea species
(Fig. 3). In 1999, only 22% of damaged flowers produced fruit compared to 52% of
undamaged, control flowers and this reduction was significant (X2dam 5 4.94, df 5 1,
178 THE AMERICAN MIDLAND NATURALIST 163(1)

FIG. 1.—The intensity of florivory in four Ipomoea species among the three study years. Values
represent the mean (SE) percentage of flowers damaged daily pooled across populations and sampling
days. Means with the same superscript in any year are not significantly different from each other, based
on a posteriori contrasts

P , 0.03). In 1999, with the native I. cordatotriloba excluded from fruit set analysis (see
Methods), there were significant differences in fruit set between the three remaining
Ipomoea species in 1999 (X2spp 5 33.47, df 5 2, P , 0.0001). A second analysis, collapsing
the three non-native species into their two pollination syndromes, reveals no difference in
fruit set between the bumble bee and two hummingbird-pollinated species (X2syn 5 0.05, df
5 1, P . 0.82) after Bonferroni correction. Thus, differences in fruit set were not explained
by pollination syndromes in 1999.
The species-by-treatment interaction was not significant in either year (1999: X2int 5 1.72,
df 5 2, P . 0.42; 2005: X2int 5 0.98, df 5 3, P . 0.81), indicating that the Ipomoea species
experienced similar reductions of fruit set when flowers were damaged.
In 2005, with all four focal Ipomoea species in the fruit set analysis, there were significant
differences in fruit set between the species (X2spp 5 28.05, df 5 3, P , 0.0001). The
differences in fruit set among the Ipomoea species was not driven by pollination syndrome
(X2syn 5 2.71, df 5 1, P . 0.10), but rather by origin. The native I. cordatotriloba had the
lowest fruit set among the four focal species, 15.0 6 3.1%, relative to the three non-native
Ipomoea, 32.9 6 2.6% (X2ori 5 3.90, df 5 1, P , 0.05), although this difference is only
marginally significant following Bonferroni correction.
Because of the way we tagged flowers in 2005, we were able to determine whether the type
of damage suffered by a flower (attractive vs. reproductive organs) influences subsequent
reproductive success. Overall, percentage fruit set in undamaged flowers was 28.0%
compared to 21.8% in flowers with damage only to the corollas, but this difference was not
significant (X2dmg 5 0.04, df 5 1, P . 0.83). However, not surprisingly, the consequences
of florivory damage to the reproductive organs were severe, relative to undamaged flowers,
2010 SOWELL & WOLFE: HERBIVORY IN IPOMOEA 179

FIG. 2.—The pattern of damage to different floral tissues in four Ipomoea species. Values represent the
percentage of damaged flowers that suffered injury to particular floral tissues. ‘Corolla’—only attractive
structures damaged; ‘Organs’—only reproductive structures damaged; ‘Flower’—both attractive and
reproductive structures damaged

FIG. 3.—The fate of damaged Ipomoea flowers. Columns represent mean 6 SE percentage of flowers
that developed into fruit
180 THE AMERICAN MIDLAND NATURALIST 163(1)

with fruit set reduced to 8.3% (X2dmg 5 7.97, df 5 1, P , 0.005), significant following
sequential Bonferroni correction.
Seed set.—The pattern of flower damage on seed production was similar to that observed
on fruit set. In 1999, florivory had a marginal effect on seed set (X2trt 5 3.39, df 5 1, P .
0.06). There were significant differences between the species for fruit set (X2spp 5 25.93, df
5 2, P , 0.0001), yet this variation in seed set was not explained by pollination syndromes
(X2syn 5 0.35, df 5 1, P . 0.55) among the three non-native Ipomoea.
In 2005, there were significant differences between the species for seed set (X2spp 5
22.34, df 5 3, P , 0.0001). Again, differences in seed set were not driven by pollination
syndrome (X2syn 5 2.95, df 5 1, P , 0.09), but rather by origin. The native Ipomoea
cordatotriloba yielded 3.2 6 0.3 seeds per fruit, whereas the non-native Ipomoea average 2.6 6
0.3 seeds per fruit (X2ori 5 3.90, df 5 1, P , 0.05), although this difference is only
marginally significant following Bonferroni correction.
There was no significant reduction in the average seeds per fruit between undamaged
flowers (1.58 6 0.14), and flowers experiencing damage only to the corolla (1.61 6 0.38;
X2trt 5 0.14, df 5 1, P . 0.70). However, seed set was approximately four times greater in
undamaged flowers (1.58 6 0.14) compared to those flowers that received damage to their
reproductive organs (0.42 6 0.23): X2trt 5 8.23, df 5 1, P , 0.005.

DISCUSSION
Flowers and floral characteristics in a variety of species have been shown to be the result of
multiple, conflicting selection pressures from pollinators and antagonists such as florivores,
nectar robbers and seed predators (Brody, 1997; Strauss, 1997; Galen, 1999; Irwin and
Brody, 1999; Pilson, 2000; Galen and Cuba, 2001). Florivores may also drive the evolution of
sexual specialization in plants (Ashman, 2000, 2002; Leege and Wolfe, 2002). Recently,
Cariveau et al. (2004) revealed that some floral traits in Castilleja linariaefolia are under
stronger selection pressure from pre-dispersal seed predators than pollinators. The
influence of florivores perhaps is not surprising, given the close association between insects
and flowering plants likely developed as an initially antagonistic relationship, due to the
increased attractiveness of flowers as food items of early angiosperm flowers relative to
gymnosperm strobili (Frame, 2003). Even though these plants likely suffered negative
effects due to loss of reproductive tissues, the reliable movement of pollen by florivores,
relative to wind pollination in gymnosperm strobili, likely led to the evolution of mutualistic
pollination system (Frame, 2003).
The main finding of this study was that all four of the Ipomoea species experienced insect-
mediated damage to their flowers, and florivory intensity was associated not with pollination
syndrome as we first hypothesized, but florivory intensity varied with whether the species was
native or introduced. The non-native, hummingbird-pollinated I. hederifolia and I. quamoclit
consistently had the lowest florivory intensity, while the native, bumble bee-pollinated I.
cordatotriloba experienced the greatest florivory intensity. In two of three study years, the
non-native, bumble bee-pollinated I. hederacea tended to have intermediate, albeit not
significantly different, florivory intensities. Florivores not utilizing I. cordatotriloba may be
more likely to host-shift onto I. hederacea, due to similar floral traits (esp. flora pigments, see
below). Florivory intensity for the four Ipomoea were fairly consistent among years, and were
comparable to florivory intensities reported for other plant species (Ackerman and
Montalvo, 1990; Bishop and Schemske, 1998; Breadmore and Kirk, 1998; Malo et al., 2001;
Leege and Wolfe, 2002). Our results support the notion that the non-native Ipomoea are
2010 SOWELL & WOLFE: HERBIVORY IN IPOMOEA 181

experiencing lower floral herbivory and that this is due to either to higher resistance to
florivores, or that florivores prefer the native I. cordatotriloba for intrinsic reasons not known.
Florivory will clearly have direct effects on fitness via male and female function, with the
most obvious impact being the loss of gametes (Louda, 1982; Bertness and Shumway, 1992;
Louda and Potvin, 1995; Lohman et al., 1996; Juenger and Bergelson, 1997; Krupnick and
Weis, 1999; Maron et al., 2002). In our study, approximately 50% of all damaged Ipomoea
flowers experienced the loss of the pistil and stamens, and this had a direct impact on
female reproductive success. In 2005, few of the flowers receiving damage to their
reproductive organs produced any fruit. Florivory may negatively impact male fitness
through loss of pollen-bearing stamens. Plants with damaged flowers should experience
reduced male reproductive success via decreased pollen export (Krupnick and Weis, 1999;
Mothershead and Marquis, 2000). Whether florivory causes pollen limitation, though, at the
community level in our Ipomoea sites is unknown (e.g., Ashman et al., 2004), though, florivory
has been shown to induce pollen limitation in other studies (Bertness and Shumway, 1992;
Cunningham, 1995; Krupnick and Weis, 1999).
Interactions with enemies that consume flowers or render flowers dysfunctional (e.g.,
disease-Alexander and Antonovics, 1988; galls-Wolfe, 1997; Wolfe and Rissler, 1999) can
impact plant reproductive success even before pollination can occur. The most obvious
influence of floral damage may be via its indirect effect on pollinator-mediated attraction.
Studies on other plant species have found reduced pollinator visitation rates when flowers
appear damaged (Johnson et al., 1995; Lohman et al., 1996; Lehtilä and Strauss, 1999;
Krupnick et al., 1999; Mothershead and Marquis, 2000; Malo et al., 2001; Canela and Sazima,
2003). This can influence male fitness if pollen removal rates are reduced and female fitness
if pollen deposition rates are reduced. In 2005, female reproductive success (fruit set) was
depressed by approximately 30% in flowers that had only their corolla damaged, but organs
left intact, suggesting that removal of attractive structures indirectly affected fruit set. Yet,
decreased attraction and reduced pollen transfer in these Ipomoea will likely not result in
complete reproductive failure since the four species are self-compatible (Ennos, 1981;
Stucky, 1985; Murcia, 1990), and can set fruit in the absence of pollinators (Wolfe and
Sowell, pers. obs.). In these Ipomoea, autogamous pollen grains may be deposited on the
stigma before or shortly after anthesis begins. Since the florivores are rarely active before
anthesis (Sowell and Wolfe, pers. obs.), damage by florivores may not occur until pollen
tubes have already passed down the style. Thus, fruit set could still occur via selfing before
the onset of floral damage. The mean number of seeds per fruit (seed set) was also not
affected by damage to the corolla, but was reduced when reproductive organs were damaged
by florivores.
Three of the four Ipomoea examined in this study are non-native and considered invasive
in the southeast U.S. (USDA, 2001). An often-cited explanation for the success of
invasiveness in the introduced range is the Enemy Release Hypothesis (Keane and Crawley,
2002) which proposes that colonization in the introduced range usually results in the escape
from enemies in the native range, allowing the introduced species to exhibit enhanced
growth and reproductive output, resulting in population expansion. We would therefore
expect enemy attack to be lower in non-native species relative to either native congeners or
the species in their native range (Maron and Vilà, 2002; Wolfe, 2002; Torchin et al., 2003;
Agrawal et al., 2005). Although the current study was not designed as a direct test of the
Enemy Release hypothesis, it is interesting that our results were consistent with expectation,
since the florivory rate was greatest on the native I. cordatotriloba compared to the non-native
Ipomoea.
182 THE AMERICAN MIDLAND NATURALIST 163(1)

It is important to point out a limitation of a comparative study such as ours that examines
multiple species. It is entirely possible that the similar patterns of florivory (or any
measurement taken on a species) between Ipomoea hederifolia and I. quamoclit is a result of the
fact that they are closely related and placed in the same subgenus (Miller et al., 1999), and
not due to the fact that they display the same pollination syndrome. Thus, the ideal study
would be one that utilizes replicate Ipomoea: (1) within subgenera that exhibit different
pollination syndromes, or (2) that exhibit the same pollination syndrome in different
subgenera. Similarly, we must be cautious to conclude that the uniqueness of I.
cordatotriloba’s greater florivory intensity is due to its being a native species since there was
no replication at the species level for native species while we did have multiple (3)
introduced Ipomoea species.
Why then, should floral herbivores exhibit a preference for the bumble bee-pollinated
Ipomoea cordatotriloba, relative to the hummingbird-pollinated Ipomoea species? It is possible
that the pattern of florivore choice is the result of a pleiotropic effect. In Ipomoea, the
production of specific plant secondary defensive compounds and specific classes of floral
pigments are not independent. Anthocyanins and secondary flavonoids are derived from
the same precursor compounds in a common biosynthetic pathway in Ipomoea purpurea, a
purple bumble bee-pollinated morning-glory (Simms and Bucher, 1996; Fineblum and
Rausher, 1997). Species with reduced anthocyanin pigmentation in their flowers, such as I.
hederifolia and I. quamoclit, may devote more precursor flavonoids to secondary defensive
compounds throughout the plant (Fineblum and Rausher, 1997), and consequently, they
may be better defended against foliar herbivory and florivory. Recent work on Nemophila
menziesii has also shown that damage to flowers can induce increased resistance to florivory
later in the season (McCall, 2006). Future work should determine if the orange and red-
flowered, hummingbird-pollinated I. hederifolia and I. quamoclit are better defended via
secondary flavonoids, relative to the blue and purple-flowered, bumble bee-pollinated
Ipomoea, and if the pattern of florivory that we observed is in part the result of a difference in
secondary flavonoids among the four Ipomoea species.

Acknowledgments.—The authors thank C. O’Neil, B. Paige, B. Penna and K. Wollett for help
conducting fieldwork; M. McAloon for identification of lepidopteran larvae; C. Barr, A. Blair, S.
Freedberg, S. Keller, M. Neiman, V. Panjeti, D. Sloan, D. Taylor and two anonymous reviewers for
valuable comments that improved the manuscript. This work was supported by grants from the National
Science Foundation (DEB-0349553) (LMW) and Georgia Southern University (Graduate College
[DRS]; College of Science and Technology Academic Excellence [DRS]; Irene Burt Boole Scholarship
[DRS]; Catalyst Grant [LMW]).

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SUBMITTED 7 JULY 2008 ACCEPTED 21 JANUARY 2009