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2011 SOUTHEASTERN NATURALIST 10(4):761–771
Susceptibility of Cultivated Native Wildflowers to Deer
Lucas W. DeGroote1,*, Holly K. Ober1, James H. Aldrich2, Jeff G. Norcini3,
and Gary W. Knox2
Abstract - Foraging preference of Odocoileus virginianus (White-tailed Deer) at ornamental
plantings was compared amongst 11 wildflower species native to north Florida and south
Georgia. Deer exhibited strong preference for Coreopsis floridana (Florida Tickseed),
C. gladiata (Coastalplain Tickseed), C. integrifolia (Fringeleaf Tickseed), and Rudbeckia
fulgida (Orange Coneflower). Browsing significantly reduced the height of Florida, Coastalplain,
and Fringeleaf Tickseeds, and reduced the number of Florida and Fringeleaf
Tickseed flowers. Browsing pressure remained high throughout the growing season; therefore,
temporary exclosures are unlikely to offer a viable solution to damage caused by deer.
Information on variation in deer preference between species and across seasons should help
private landowners and public land managers make strategic decisions regarding which
species to establish at residences, food plots, or roadside beautification projects.
Odocoileus virginianus Zimmermann (White-tailed Deer) are capable of
causing extensive damage to ornamental plants and agricultural crops (Conover
1997, Conover and Kania 1988, Garrison and Lewis 1987, Stratton and Smathers
1996). Conversion of natural habitat to agricultural lands and housing developments
coupled with restriction of hunting in developed areas has increased both
the contact between humans and deer as well as the frequency and extent of damage
inflicted on ornamentals and crops by deer throughout the southeastern US
(Harden et al. 2005).
Deer can be deterred from browsing ornamentals and crops through a variety
of options; however, nearly all of these options are costly, unsightly, work for
only a brief period of time, or are considered objectionable to some members of
the public (Andelt et al. 1994, Conover 2001, Mulinas et al. 1994, Rosenberry
et al. 2001). A more viable, long-term option for preventing deer damage to
ornamentals and crops entails selecting species or varieties that deer find unpalatable
for most of the calendar year (Conover and Kania 1988). Because deer
preferences vary seasonally (Garrison and Gedir 2006, Godvik et al. 2009), an
understanding of the changes in deer foraging habits throughout the growing season
would allow homeowners to restrict efforts to protect ornamentals to those
periods when the plants are most susceptible to deer browsing.
1Department of Wildlife Ecology and Conservation, North Florida Research and Education
Center, University of Florida, 155 Research Road, Quincy, fl32351. 2Department
of Environmental Horticulture, North Florida Research and Education Center, University
of Florida, 155 Research Road, Quincy, fl32351. 3OecoHort, LLC, 726 Riggins Road,
Tallahassee, fl32308. *Corresponding author - firstname.lastname@example.org.
762 Southeastern Naturalist Vol. 10, No. 4
Another potential concern regarding deer foraging on ornamentals involves the
possible attraction of deer to roadside plantings. Many states have adopted roadside
beautification projects such as wildflower planting right-of-way programs.
The Florida Wildflower Foundation (2011) and the Georgia Department of Transportation
(GDOT 2011) offer grants to promote highway beautification through
landscape programs, with the hope that these programs will lower maintenance
costs, reduce erosion, and increase driver alertness. While we are not aware of any
research investigating the relationship between highway beautification projects
and deer-vehicle collisions, alteration of roadside vegetation is recognized by
wildlife biologists and department of transportation administrators as one of the
more cost-effective options for mitigating deer-vehicle collisions (Sullivan and
Messmer 2003). It therefore seems logical that planting native species found to be
desirable or undesirable by deer could increase or decrease, respectively, the frequency
of deer-vehicle collisions. Understanding which species of wildflowers are
most susceptible to deer damage would be helpful in providing recommendations
regarding which species should not be planted along roadsides.
Many studies that have attempted to determine preferences of foraging deer
have been conducted under artificial conditions with captive deer that may not
feed in a manner typical of free-ranging individuals (Crouch 1966, Pepin et al.
2006, Radwan and Crouch 1974, Sauve and Cote 2006). Assessing the preferences
of free-ranging deer whose behavior has not been altered in any way is
a superior means of evaluating impacts of browsing pressure on ornamentals,
particularly if conducted in areas with naturally high densities of deer.
We developed a project to investigate foraging preference of wild Whitetailed
Deer among annual and perennial Asteraceae native to north Florida and
south Georgia. Several species of Coreopsis were chosen for our study because
we had previously observed a difference in deer preference amongst species
of this genus, and because Coreopsis (undesignated species) is Florida’s state
wildflower and is therefore desirable in natural, roadside and ornamental plantings.
The 11 species selected for this study, despite their relatedness, represent
wildflowers with both upland and wetland habitat preferences; spring, summer,
fall or continuous bloom periods; annual, short-lived perennial, and perennial
characteristics; as well as rhizomatous, stoloniferous, and crown growth types.
The objectives of our study were to determine (1) which wildflower species
are most preferred by foraging deer in this region, (2) the effect of browsing on
growth and flower production, and (3) if any species would benefit from temporary
protection against foraging deer during a particular period during the year.
Our results should ultimately reduce economic losses incurred by individuals
interested in maintaining ornamental plantings and potentially decrease the likelihood
of deer-vehicle collisions in areas with purposeful roadside plantings.
We initiated a study to evaluate foraging preference of free-ranging White-tailed
Deer for native wildflowers at the North Florida Research and Education Center
2011 L.W. DeGroote, H.K. Ober, J. Aldrich, J.G. Norcini, and G.W. Knox 763
(NFREC) in Quincy, fl(30°32'42"N, 84°35'39"W) from April to November of
2008 and 2009. Although the exact density of deer at our study area is not known,
deer density in northern Gadsden County is estimated to be among the highest in
the state of Florida (QDMA 2011). Preference was tested amongst 11 species of
native annuals and perennials (Asteraceae) selected for their availability (seeds
collected <100 mi from study area), potential usefulness in landscape settings,
and anecdotal observation of deer browsing preference. Species tested included
Coreopsis basalis Blake (Goldenmane Tickseed), C. floridana E.B. Sm. (Florida
Tickseed), C. gladiata Walt. (Coastalplain Tickseed), C. integrifolia Pior. (Fringeleaf
Tickseed), C. lanceolata L. (Lanceleaf Tickseed), C. leavenworthii T. and
G. (Leavenworth’s Tickseed), Gaillardia pulchella Foug. (Firewheel), Ratibida
pinnata Vent. (Yellow Coneflower), Rudbeckia fulgida Ait. (Orange Coneflower),
R. hirta L. (Black-eyed Susan), and R. mollis Ell. (Softhair Coneflower). Testing
was performed at 2 experimental plots, 650 m2 and 850 m2 in size. Plot 1 was
located near a busy roadway (50 m), pond (50 m), deciduous forest (50 m), and
fields of peanuts and soybeans (<300 m), while plot 2 was located near private
property with coniferous trees but no understory (50 m), mixed coniferous-deciduous
forest (160 m), fields of winter grasses (50 m), and vegetable plantings
(<400 m; pumpkins, squash, green peppers, and onions). Each plot was covered
in landscape fabric, divided into 8 replicated blocks, and planted each year with
3 plants per species in each block. To reduce competition, facilitate growth, and
provide equal opportunity for browsing by deer, container-raised seedlings were
planted at least 0.6 m apart, fertilized when planted (5 g of Osmocote 15-9-12,
12–14 month southern formulation), irrigated during drought periods until established,
and hand-weeded as needed. Three of the 8 blocks in each plot were
protected from deer with a tall fence, and 5 were left unprotected. Plant height,
number of flowers (0 – 1, 2 – 5, 6 – 20, or >20), and deer damage (presence or
absence of browsing) were recorded every 2 weeks from 7 April (shortly after
planting) through the end of their effective growing season, 17 November.
We quantified deer preference and the effect of deer on wildflower species in
4 ways: percentage of plants browsed, effect of browsing on flower production,
severity of browse damage, and timing of browse damage. All statistical analyses
were conducted with the program R version 2.8.1 (R Development Core
The percentage of plants browsed was compared amongst the 11 species
using a general linear model (GLM) with quasibinomial errors (Crawely
2007). Unlike traditional contingency tables, the aforementioned GLM is
more robust for datasets with an unbalanced design due to missing values (in
our case, from plant death). Plants that died before browsing occurred were
excluded from the analysis, and a plant was considered browsed if it was
browsed at any time throughout the year. To determine if browsing differed
between years and plots, we modeled the ratio of browsed to unbrowsed plants
as a binomial denominator against species (a multilevel factor), year, plot, and
all two-way interactions. We used an iterative process whereby whichever
independent variable (plot, year, species, or the interaction variables) that
764 Southeastern Naturalist Vol. 10, No. 4
explained the least amount of variability was removed from the full model,
and P-values were calculated from a likelihood ratio test comparing the reduced
model to the full model. This process was repeated until the likelihood
ratio was significantly different, indicating the most parsimonious model had
been identified (Diggle et al. 2002). We then used the same model-reduction
process to identify which species were browsed more frequently. Species were
removed one at a time from the aforementioned most parsimonious model
and compared to the prior model. Resulting P-values were bootstrapped to
calculate q-values, the analogous Bayesian posterior P-value, to account for
positive false discovery rates (pFDR) resulting from multiple comparisons
(Dabney and Storey 2010, Storey 2003).
We compared flower production between browsed and unbrowsed plants for
the four most commonly browsed species using an analysis of variance (ANOVA)
and Tukey’s honest significant differences (Tukey’s HSD). Because plants inside
and outside of the deer enclosure provided useful information on flower performance,
all plants were included in the analysis. We created a single ANOVA
model to compare the maximum number of flowers per plant from any sampling
period (using category midpoints of 0.5, 3.5, 12.5, and 25) by species, year, plot,
browse damage (presence or absence), and their 2-way interaction variables.
We compared the severity of damage amongst the four most commonly
browsed species by quantifying height lost from browsing. Height loss was
calculated by taking the height of a plant when browsing was observed and subtracting
it from the height measured 2 weeks prior. If an individual plant was
browsed more than once throughout the study, height loss was averaged to avoid
pseudo-replication. To account for heterogeneity present across species and year,
we used a general least squares model to determine average height loss for each
Lastly, we used generalized linear mixed models with binomial errors to investigate
browse timing for the four most commonly browsed species. The full
model consisted of presence or absence of browse damage regressed by plot,
year, species, and the first- through third-order polynomial of sampling week
(mean centered). Each plant was included as a random effect to account for
repeated measurements of the same plants over time. The most parsimonious
model was identified with the same iterative process used in the percent browsed
analysis. Browse timing was graphically represented by locally weighted least
squares (Lowess) curves showing the percentage of browsed plants over the year
for each species-year combination.
The percentage of plants browsed by deer was significantly higher for Florida
Tickseed, Coastalplain Tickseed, Fringeleaf Tickseed, and Orange Coneflower
than for Lanceleaf Tickseed, Goldenmane Tickseed, Leavenworth’s Tickseed,
Firewheel, Yellow Coneflower, Black-eyed Susan, and Softhair Coneflower
(q-value ≤ 0.05, Table 1). The interaction between plot and species was nonsignifi
cant (likelihood ratio test: P = 0.12), and the most parsimonious model
2011 L.W. DeGroote, H.K. Ober, J. Aldrich, J.G. Norcini, and G.W. Knox 765
indicated that the percentage of browsed plants was significantly greater at plot
1 than plot 2 (P < 0.01).
Deer browsing had a significant effect on maximum flower production (P <
0.01, F1 , 317 = 28.65), where browsed plants produced fewer flowers per plant
(mean = 6.4) than unbrowsed plants (mean = 11.7; Fig.1). Tukey’s HSD revealed
Figure 1. Maximum number of flowers produced in the presence (white) or absence (gray)
of browse damage for the four most frequently browsed species of wildflowers. Bars represent
standard errors and significant differences are indicated by * (P ≤ 0.05).
Table 1. Preference of White-tailed Deer for native wildflower species based on results of general
linear model analyses. The q-value is the analogous P-value for multiple comparisons positive false
discovery rates. Significant values and percentages are shown in bold.
Species q-value Percent browsed
Coreopsis lanceolata (Lanceleaf Tickseed) 0.42 3%
Ratibida pinnata (Yellow Coneflower) 0.42 3%
Coreopsis basalis (Goldenmane Tickseed) 0.42 5%
Gaillardia pulchella (Firewheel) 0.16 5%
Rudbeckia mollis (Softhair Coneflower) 0.18 17%
Rudbeckia hirta (Black-eyed Susan) 0.16 23%
Coreopsis leavenworthii (Leavenworth’s Tickseed) 0.16 27%
Rudbeckia fulgida (Orange Coneflower) 0.04 42%
Coreopsis gladiata (Coastalplain Tickseed) 0.03 48%
Coreopsis floridana (Florida Tickseed) 0.01 60%
Coreopsis integrifolia (Fringeleaf Tickseed) <0.01 67%
766 Southeastern Naturalist Vol. 10, No. 4
that browsed Florida Tickseeds produced 49% fewer flowers (P = 0.01) than
unbrowsed plants (mean difference = 5.98), while browsed Fringeleaf Tickseeds
produced 58% fewer flowers (P < 0.01) than unbrowsed plants (mean difference
= 11.03). Browsing had no effect on Coastalplain Tickseed or Orange Coneflower
flower production (P = 0.99).
Because browsing at plot 2 was infrequent, the severity and timing of
browse damage was investigated at plot 1 only. Height of Florida, Coastalplain,
and Fringeleaf Tickseed was significantly reduced by browsing (P <
0.01 for 2008 and 2009; Fig. 2), while Orange Coneflower height increased
from growth faster than height was reduced by browsing (P < 0.01 for 2008
Browsing began 5 weeks earlier (P < 0.01) in 2009 (2 June) than 2008
(5 July). In both years, the percentage of browsed Florida and Coastalplain
Tickseed remained high (>50%) from the initial date of browsing to the end
of the growing season (Fig. 3). In 2008, Fringeleaf Tickseed also had a high
percentage of plants browsed from the initial date of browsing onward, but the
percentage of plants browsed in 2009 was only high from 2 June to 10 August
(P < 0.01). The percentage of browsed Orange Coneflower was high (>50%)
from 2 June to 10 September 2009 only (P < 0.01).
Figure 2. Average height loss of the three species of wildflowers browsed most heavily by
White-tailed Deer (height prior to damage – height after damage) at plot 1. Height loss in
2008 is represented by gray bars, 2009 is in white. Bars represent standard errors.
2011 L.W. DeGroote, H.K. Ober, J. Aldrich, J.G. Norcini, and G.W. Knox 767
Four of the 11 native wildflower species were more susceptible to browsing
by White-tailed Deer: Florida, Coastalplain, and Fringeleaf Tickseed, as well as
Orange Coneflower (Table 1). Research at other sites has shown that browsing by
White-tailed Deer can inhibit wildflower growth and flower production, altering
the structure and dynamics of plant populations in natural settings (Augustine
and DeCalesta 2003, Barrett and Stiling 2006, Frankland and Nelson 2003,
Kettenring et al. 2009, Rooney 2001, Rooney and Gross 2003). For example,
browsing increased mortality, reduced plant height, inhibited flower production,
and decreased fecundity of Trillium grandiflorum (Michx.) Salisb. (White Trillium)
in North American forests (Frankland and Nelson 2003, Rooney and Gross
2003). Closer to our study area, an experimental study in central Florida demonstrated
that Liatris ohlingerae (S.F. Blake) B.L. Rob. (Florida Blazing Star) were
shorter, less likely to flower, and had fewer inflorescences in areas where deer
were not excluded (Kettingring et al. 2009). We found that Orange Coneflower
Figure 3. Lowess curves representing the percentage of browsed plants over time (April
to November) at plot 1 for the four preferred species: (A) Florida Tickseed, (B) Coastalplain
Tickseed, (C) Fingeleaf Tickseed, and (D) Orange Coneflower. Solid line and points
correspond to browsing in 2008, dashed lines and open circles correspond to 2009. Tick
marks on the x-axis indicate the first sampling period of each month. Three sampling
periods occurred in July, two in every other month.
768 Southeastern Naturalist Vol. 10, No. 4
was the only species of the four most heavily browsed species (i.e., species for
which >40% of plants were damaged) that was able to regrow and flower to
sustain an attractive appearance (Figs. 1 and 2). Florida and Fringeleaf Tickseed
plants browsed by deer produced fewer flowers and often perished (65% and
25% mortality, respectively). We did not observe any effect of browsing on the
maximum number of flowers produced by Coastalplain Tickseed because most
plants of this species did not survive to flower (17% survival for unbrowsed,
13% for browsed). Coastalplain Tickseed’s low survival rate is likely explained
by the relatively dry conditions in our ornamental landscape setting compared
with the species’ preference for wetter habitats (Norcini and Aldrich 2007).
Of the few Coastalplain Tickseeds which survived to flower (n = 15), browsed
plants produced fewer flowers (mean = 9.6) than unbrowsed individuals (mean =
18.0). Because of their overall susceptibility to browsing, cultivation of Florida,
Coastalplain, and Fringeleaf Tickseeds for aesthetic purposes in north Florida
and south Georgia may be extremely difficult without protection from deer (i.e.,
exclusion fences; Rosenberry et al. 2001). Unprotected, long-term ornamental
plantings of these annual and short-lived perennial species would likely require
re-seeding to compensate for reduced seed production brought about by browsing.
Conversely, the relative unpalatability of Goldenmane Tickseed, Lanceleaf
Tickseed, Leavenworth’s Tickseed, Firewheel, Yellow Coneflower, Black-eyed
Susan, and Softhair Coneflower, and the ability of Orange Coneflower to withstand
browsing, could make them desirable species for nurseries and ornamental
gardens in areas with high deer densities.
The strong preference White-tailed Deer exhibited for Florida, Coastalplain,
and Fringeleaf Tickseeds may attract deer to areas with purposeful plantings
of these species, an effect which could be desirable or undesirable. Planting
these species in landscape beds could attract deer to properties for wildlife
viewing or hunting. Food plots, plantings intentionally established by hunters
to concentrate deer in particular areas, often consist of non-native plants or agriculture
crops which may require annual planting, herbicide, insecticide, and
fertilizer (i.e., Hehman and Fulbright 1997, Johnson and Dancak 1993). Using
native wildflowers adapted to north Florida and south Georgia that are known
to be favored by deer and also known to be capable of sustaining browsing
could reduce costs and environmental impacts typically associated with foodplot
establishment and maintenance.
Browsing at our study site began late summer in 2008 (28 July) and early
summer the following year (2 June 2009). The intensity of browsing over time
also varied between years and across species (Fig. 2). For example, Florida,
Coastalplain, and Fringeleaf Tickseed were intensely browsed from late July to
mid-November in 2009, while the intensity of browsing on Florida and Fringeleaf
Tickseeds was lower overall in 2008 and slacked for 6 weeks between late
August and early October. The time of year in which browsing began and the
intensity of browsing may have differed between years because food preferences
of free-ranging deer can be influenced by the availability of local food resources
(Garrison and Gedir 2006) and differential risk of predation (Brown and Kotler
2011 L.W. DeGroote, H.K. Ober, J. Aldrich, J.G. Norcini, and G.W. Knox 769
2004, Godvik et al. 2009). Likewise, food availability or predation risk could
also explain why damage was significantly different between our two study
plots. Specifically, plot 1 was located closer to agricultural fields with crops deer
find highly palatable (peanuts and soybeans), while plot 2, being located farther
away from roads, water, and deciduous cover, may have posed a higher risk for
predation. Because of the variability we observed between plots and years, we
recommend that private land owners and nurseries monitor local deer damage to
Florida, Coastalplain, or Fringeleaf Tickseed to assess the cost versus benefit of
protecting these species throughout the growing season.
Our results indicate that White-tailed Deer preference for wildflowers changed
over time and differed amongst species, which suggests that carefully planned
highway beautification projects could reduce the risk of deer-vehicle collisions.
Traditional roadside plantings in the southeastern US consist of clover and cover
grass which can be palatable for deer year-round (Chapman et al. 2009, Murphy
et al. 1985). Because deer in our study did not browse any wildflowers during the
spring, replacing traditional roadside plantings with native wildflower plantings
that flower during this time of year could potentially reduce deer-vehicle collisions.
Moreover, planting other species which we found to be unpalatable for deer
year-round could further reduce this risk. However, a multi-year, multi-location
study comparing traditional roadside plantings to native grass-wildflower stands
would be needed to elucidate the effect of plantings on deer-vehicle collisions.
The four most commonly browsed species all flowered in the late summer
or fall. None of the spring or early summer blooming wildflowers in our study
were browsed. This pattern may have arisen because deer find wildflower plants
in bloom less palatable or because other highly preferred foods were more available
early in the year. In comparison to the other species tested, the four preferred
species are naturally found in wetter habitats (Clewell 1985) and possess more
succulent stems and leaves (Norcini and Aldrich 2007). Taken together, our results
indicate that White-tailed Deer in north Florida may prefer wildflowers that
favor wet habitats, possess succulent foliage, and/or bloom in the late summer or
fall; however, further study is needed to determine if these preferences remain
true across a wider diversity of habitats and native wildflower species.
We thank T. Batey, A. Brock, J. Crowell, and S. Wright for their assistance establishing
and maintaining research plots, planting and irrigating wildflowers, collecting data,
and entering data.
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