Garden Plants Attractiveness To Bees
Garden Plants Attractiveness To Bees
Garden Plants Attractiveness To Bees
12178
Summary
1. Pollinating insects are globally declining, with one of the main causes being the loss of flow-
ers. With the value of countryside reducing, urban areas, particularly gardens, are increasingly
recognized as of benefit to wildlife, including flower-visiting insects.
2. Many gardeners specifically select plant varieties attractive to wildlife. Given the wide public
interest, many lists of recommended varieties have been produced by both amateurs and
professional organizations, but appear not to be well grounded in empirical data. These lists,
however, are not without merit and are an obvious starting point. There is clearly a need to
put the process onto a firmer footing based more on data and less on opinion and general
experience.
3. We collected data over two summers by counting flower-visiting insects as they foraged on
32 popular summer-flowering garden plant varieties in a specially planted experimental garden,
with two smaller additional gardens set up in year two to check the generality of the results.
With many thousands of plant varieties available to gardeners in the United Kingdom, and
other countries or regions, it would have been an impossible task to make a comprehensive
survey resulting in a complete and authoritative list.
4. Our results are valuable and encouraging. Garden flowers attractive to the human eye vary
enormously, approximately 100-fold, in their attractiveness to insects. Insects, especially bees
and hover flies, can be attracted in large numbers with clear differences in the distribution of
types attracted by different varieties.
5. Our results clearly show that there is a great scope for making gardens and parks more bee-
and insect-friendly by plant selection. Horticulturally modified plant varieties created by plant
breeding, including hybrids, are not necessarily less attractive to insects and in some cases are
more attractive than their wild-type counterparts. Importantly, all the plants we compared
were considered highly attractive to humans, given that they are widely sold as ornamental
garden plants.
6. Helping insect pollinators in gardens does not involve extra cost or gardening effort, or loss
of aesthetic attractiveness. Furthermore, the methods of quantifying insect-friendliness of plant
varieties trialled in this study are relatively simple and can form the basis of further research,
including ‘citizen science’.
green spaces, such as parks and gardens (Helden & authoritative list. What our data do show, however, is
Leather 2004; Matteson, Ascher & Langellotto 2008; Owen valuable and encouraging. Garden flowers attractive to the
2010), with private gardens often being the largest and human eye vary enormously, approximately 100-fold, in
probably the most important component (Goddard, Dougill their attractiveness to insects. This shows that plant selec-
& Benton 2010). In the United Kingdom, 87% of house- tion can make a great difference in the value of gardens
holds are associated with a garden (Davies et al. 2009) and and parks to flower-visiting insects, and at no additional
gardening is a popular hobby (Taylor 2002). In addition, cost. Insects, and especially bees, can be attracted in large
many gardeners are supportive of wildlife, with most UK numbers with clear differences in the distribution of types
gardeners (74–78%) engaging in some form of ‘wildlife attracted by different garden plant varieties.
gardening’. That is, doing something to attract or encour-
age wildlife (Good 2000), including the 31% who select
plants attractive to wildlife or the 66% who feed birds in Materials and methods
their garden (Mew et al. 2003; DEFRA 2007).
Garden plants are often non-native, and this may reduce EXPERIMENTAL PLANT VARIETIES AND FLOWER BEDS
their usefulness to some wildlife. For example, many We studied 32 garden plant varieties that include 19 species and
herbivorous insects have a narrow range of suitable food hybrids, both native and exotic to Britain, with particular focus
plants (Novotny & Basset 2005; Dyer et al. 2007). How- on varieties of lavender (Lavandula spp.), as it is known to be
ever, this does not prevent them from being useful to attractive to bees (Pawelek et al. 2009; for full list see Table S1,
Supporting information). Varieties were selected based on the
flower-visiting insects seeking nectar and pollen, as these
following three criteria: they were (i) popular as garden plants in
are general resources. Nectar, for example, is mainly sugar their own right due to their attractive flowers or foliage (e.g.
and water (Nicolson & Thornburg 2007), and so it is Lamb’s ear, Stachys byzantina), (ii) widely and easily available for
edible whether from a native or a non-native plant. Many purchase and (iii) flowered mainly or exclusively in late summer,
garden plants have also been bred to alter their appear- July and August, as these are the months when honeybee foraging
distances in the same area are greatest (M.J. Couvillon, R.
ance, such as by the ‘doubling’ of petals, which may reduce
Sch€urch & F.L.W. Ratnieks, unpublished data), indicating chal-
floral rewards or their accessibility (Comba et al. 1999; lenging foraging conditions and, therefore, the period when gar-
Corbet et al. 2001). den flowers can be particularly beneficial to flower-visiting insects.
Given the public interest in helping wildlife, a large The main experimental flower bed was on the University of
number of recommended plant lists have been produced, Sussex campus (lat: 50865646, long: 0090771943) on chalky soil
of the South Downs. All 32 varieties were planted in 1 9 1 m
by both amateurs (e.g. Baines (2000); Lavelle & Lavelle
patches, two patches per variety, in two concentric circles (inner
(2007)) and professional organizations (e.g. Royal Horti- diameter 122 m, outer 192 m), with one variety per circle in a
cultural Society (2011); Xerces Society (2011)). However, random position (Fig. 1). There were gaps of c. 05 m (inner) and
these appear not to be well grounded in empirical data. 10 m (outer) between adjacent patches within the same circle and
For example, Thompson (2006) referred to one list of wild- 15 m between the circles. This arrangement was chosen to elimi-
nate any edge effects, which might have affected insect visitation.
life friendly plants produced by Natural England, a
Data were collected in both 2011 and 2012.
government-funded agency responsible for protection and Additionally, to ensure our results were not location specific, for
improvement of the natural environment, as ‘looks very example, due to local soil conditions or insect abundance, 13 of the
much as if it was put together late one Friday afternoon’. 32 varieties (Table S1) were planted at two additional locations in
In addition, lists of bee- and butterfly-friendly plants vary 2012. This subset was chosen to confirm certain notable trends seen
in the data at the end of the first season (2011). For example, Bora-
greatly even when they are for the same country, suggest-
go was mostly visited by honeybees, while Lavandula mostly by
ing that the underlying information is based mainly on bumblebees. L. 9 intermedia received more insect visits than
personal observations, experience, opinion and, perhaps, L. angustifolia, while sharp colour contrast (traditional blue/purple
uncritical recycling of earlier lists (M. Garbuzov & F.L.W. vs. white) had no effect on the number of visits. Open-flowered
Ratnieks, unpublished data). Dahlia varieties were more attractive than those with more modi-
fied flower forms. Origanum and Stachys seemed to have dispro-
Lists of bee- and butterfly-friendly plants are not with-
portionately large numbers of visits by ‘other’ wild bees, while
out merit and are an obvious starting point for determin- Erysimum was most attractive for butterflies and moths. One loca-
ing which plants are good for flower-visiting insects. tion was 45 km away at Plumpton College (lat: 50905665, long:
However, there is a need to put the process onto a firmer 0074753791) where the soil is also chalky, and the other 263 km
footing based more on data and less on opinion and gen- away in FR’s private garden in Magham Down (lat: 50880426,
long: 028488247), where the soil is sandy. Only one 1 9 1 m patch
eral experience. This study is an attempt to do this. We
per variety was planted, and the patches were arranged in a line
collected data over two summers in which flower-visiting with 30 cm gaps.
insects were counted as they foraged on 32 popular garden Perennials were bought in pots from nurseries and garden
plant varieties in a specially planted experimental garden. centres and planted in June 2011 (University of Sussex) and May-
In addition, two smaller gardens were set up in year two June 2012 (Plumpton College & Magham Down). Borage (Borago
officinalis), which is an annual, was sown in May each year to
to check the generality of the results. With many thou-
give peak flowering in July–August. Viper’s bugloss (Echium
sands of plant varieties available to gardeners in the Uni- vulgare) is a biennial, flowering in the second year of its life cycle.
ted Kingdom, it would have been an impossible task to However, we were able to induce flowering in the first year
make a comprehensive survey resulting in a complete and by keeping young seedlings in a greenhouse at 24:0 light/dark
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
Garden plants for flower-visiting insects 3
32
On each day of data collection, the bloom intensity of patches
N 27
17 7
16
(a) was quantified by assigning a score 0 (absence of bloom), 1 (<⅓ of
22 23
maximum), 2 (⅓–⅔ of maximum) or 3 (full bloom, >⅔ of maxi-
26 9 12 22 21
2 30
4
mum; after Anderson & Hubricht 1940) and included as a
1
1 9 covariate in the analyses. In addition, as corolla length is known
14 28
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
4 M. Garbuzov & F. L. W. Ratnieks
Table 1. Breakdown of main insect groups that were grouped together in the analyses
Bombus subgroups follow Fussell & Corbet (1992). For the relative abundance of groups see Figs 2 and S2.
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
Garden plants for flower-visiting insects 5
32%
5% COMPARISON OF LAVENDER VARIETIES
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
6 M. Garbuzov & F. L. W. Ratnieks
8 c c c
d d d d
7 e e e e e e e
6 f f
g g g
5 h h h h h h
4 i i i
j j j j j j j j
3 k k k k
2 l l l l l l l l l l l
m m m m m m m m m m m m m m m
1
0
a a a a a (b) 2012
b b b b b b
Daily mean insects per snapshot per m 2
8
c c c c c c c c c
7 d d d d
e e
6
f f f f f f f f f f f f f f f f f f f f f f
5 g g g g g g g g g g g g g g g g g g g g g g
h h h h h
4
3
2
1
0
Fig. 3. Daily mean numbers of insects per snapshot per 1 9 1 m patch recorded on 32 garden plant varieties at the University of Sussex
in 2011 (a) and 2012 (b), Letters above bars represent significant differences based on Tukey’s HSD test, where varieties sharing a common
letter are not significantly different from each other at a = 005. Full plant names are given in Table S1.
non-significant [honeybees vs. other bees (r = 0169, the 32 plant varieties at the University of Sussex in the
P = 0354), honeybees vs. hover flies (r = 0251, P = 0165), total number of insects attracted. This clearly shows that
honeybees vs. butterflies & moths (r = 0157, P = 0390), there is great scope for making gardens and parks more
bumblebees vs. other bees (r = 0077, P = 0675), bumble- bee- and insect-friendly by judicious plant selection.
bees vs. hover flies (r = 0266, P = 0141)] (Fig. 7a–c). The Importantly, this need not involve extra cost or gardening
significant correlations (Figs 6b and 7c) remained signifi- effort, or, indeed, a loss of aesthetic attractiveness, given
cant after Bonferroni correction (a = 005/8). that all the plants we compared were considered to be
highly attractive, and were easily available at comparable
and low prices. Our results can be considered as a contri-
Discussion
bution to the lists of recommended garden plants.
The results showed very large, approximately 100-fold However, this should be done with caution, as we only
(c. 300-fold in 2011, c. 80-fold in 2012), variation among compared 32 varieties, which is a very small proportion of
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
Garden plants for flower-visiting insects 7
4
were bumblebees plus a small percentage of other bee gen-
era. Hover flies were always the next most abundant taxon
(7–10%). Butterflies and moths (1–3%) and all other
2 insects (1–3%) were always a small percentage. Overall,
our results suggest that garden plants can easily help bees,
which showed up in large numbers, by providing forage.
0
This agrees with Goulson et al. (2010), who found evi-
Mean number of insects per snapshot per m2 at
4
P = 0·002 and especially honeybees [maximum foraging range c.
rs = 0·791 10 km, (Beekman & Ratnieks 2000)] can forage at long
distances from their nest and thus are able to exploit gar-
den resources. By contrast, butterflies and moths, being
2 relatively scarce garden flower-visitors, can perhaps not be
as easily helped by garden flowers, despite not being cen-
tral place foragers. There is also little evidence that the
0
abundance of adult resources, apart from shelter, has any
0 2 4 6 8 impact on population size or trends in European butterflies
Mean number of insects per snapshot per m2 at (Thomas, Simcox & Hovestadt 2011).
the University of Sussex flower bed in 2012
The absence of a positive correlation between the attrac-
Fig. 4. Correlations between the number of insects per snapshot tiveness of plant varieties to honeybees and bumblebees
recorded on different plant varieties at the University of Sussex (Fig. 6a) suggests that their foraging preferences do not,
flower bed in 2012 with that in (a) 2011, and (b) the additional generally, coincide. Furthermore, the absence of a negative
flower beds at Plumpton College (●) and Magham Down (○) in correlation seems to suggest that these bees do not appear
2012.
to be in competition with each other. However, N.J.
Balfour, S. Gandy & F.L.W. Ratnieks (unpublished data)
the thousands of varieties available (Cubey & Merrick showed competition on L. 9 intermedia ‘Grosso’ experi-
2011) with similar habit (small shrubs and herbaceous mentally. In particular, honeybee numbers increased c. 30
plants suitable for a mixed border). fold on patches from which bumblebees were excluded
Attractiveness of plant varieties correlated strongly (Fig. 6a). It is likely, therefore, that the lack of correlation
between 2011 and 2012 at the University of Sussex between honeybees and bumblebees reflects both the effects
(Fig. 4a). It also correlated between the University of of preferences and competition. As these two types of bees
Sussex and the two additional flower beds at Plumpton were the most abundant flower-visitors, each probably has
College and Magham Down (Fig. 4b). This shows that the capacity to affect the other via consumptive competi-
our results apply generally to a wider area and are not tion (N.J. Balfour, S. Gandy & F.L.W. Ratnieks unpub-
unduly year- or location-specific. Some variation among lished data). In the case of L. 9 intermedia ‘Grosso’, the
locations may be due to local conditions (e.g. microclimate mean corolla tube length of 72 mm was experimentally
or soil type) or differences in the flower-visiting insect com- shown to disadvantage honeybees (mean tongue length
munities present. However, most insect species or groups 66 mm) vs. bumblebees (mean tongue length 78 mm) by
we recorded are common, so would be present in almost causing longer flower-handling times (Balfour, Garbuzov
any area, but not necessarily in the same proportions. Sim- & Ratnieks 2013).
ilarly, some variation between the 2 years could be driven Plant variety attractiveness was similar between the
by annual fluctuations in insect populations. Additionally, short-tongued (B. terrestris/lucorum group) and the long-
in our study, the variation observed between years could tongued (B. hortorum and B. pascuorum groups) bumble-
be due to the different stages of establishment of perennial bees, perhaps, reflecting preferences common to Bombus in
plant varieties. In 2011, the plants had been put into the general (Fig. 6b) or the fact that tongue length variation
patches soon after being delivered from suppliers, who among bumblebees had little effect in our gardens, despite
grew them in pots, while in 2012, most varieties had had reported effects being noted in the literature (Goulson
an extra year in the ground to establish. et al. 2005; Goulson, Lye & Darvill 2008). Nepeta 9
Although variation in relative abundance of insects faassenii ‘Six Hills Giant’ stood out from this correlation,
may explain a small proportion of variation in plant being very attractive to long-tongued bumble bees [length
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
8 M. Garbuzov & F. L. W. Ratnieks
d
5 (a)
Mean number of insects per snapshot per m2
Honeybees
cd
2-banded bumblebees
4 3-banded bumblebees abc abcd abc bcd
Brown bumblebees
Other bumblebees
3 Other bees
Hover flies
Butterflies & moths
Other insects
2
a ab ab a ab a a
1
0
Arctic Folgate Hidcote Imperial Melissa Rosea Edelweiss Gros Grosso Hidcote Old Sussex Anouk
Snow Gem Lilac Bleu Giant English
100
(b) Fig. 5. (a) Numbers of insects per snapshot
Bloom duration in 2012 (days)
0
Arctic Folgate Hidcote Imperial Melissa Rosea Edelweiss Gros Grosso Hidcote Old Sussex
*
Anouk
out when plants were well established, thus
showing natural phenology. * L. stoechas
Snow Gem Lilac Bleu Giant English ‘Anouk’ was in poor condition and did not
Lavandula angustifolia Lavandula × intermedia Lavandula reach full bloom in 2012. Photographs
stoechas courtesy Dr. Simon Charlesworth.
85–125 mm (Goulson et al. 2005)], but relatively choices based on nectar and pollen rewards in bees (Seeley
unattractive to short-tongued species (length 75–76 mm), 1995; Goulson & Osborne 2010) and hover flies (Haslett
possibly due to its relatively long corolla tube 1989) and nectar rewards in other insects (Kim, Gilet &
(119 02 mm). However, other plant species with simi- Bush 2011). Our data showed no effect of corolla tube
larly long corolla tubes were attractive to short-tongued length (Table S2). However, in specific cases, corolla tube
species due to large corolla width (e.g. E. vulgare), which length may be important. In the case of lamb’s ear (Sta-
allowed short-tongued insects to place their whole head or chys byzantina), its attractiveness to wool-carder bees (An-
body far into the flower, reducing or eliminating the need thidium manicatum) is probably due to the abundant leaf
for a long tongue. In general, attractiveness did not corre- trichomes (pubescence) and possibly trichome secretions,
late between honeybees and bumblebees on the one hand, which are collected by females as nest lining material
and other bees, hover flies and butterflies + moths on the (M€ uller, Topfl & Amiet 1996; Payne, Schildroth & Starks
other, with the exception of the positive correlation 2011). In addition, lamb’s ear flowers are also visited by
between bumblebees vs. butterflies + moths (Fig. 7). How- wool carders. Some plants may be more attractive than
ever, certain plants stood out as particularly good for others by virtue of their longer flowering period. For
other, non-Apis and non-Bombus, bees (Origanum vulgare, example, N. 9 faassenii ‘Six Hills Giant’ and Erysimum li-
E. vulgare, S. byzantina, Achillea millefolium), hover flies nifolium ‘Bowles Mauve’, which are sterile hybrids unable
(O. vulgare, A. foeniculum, E. vulgare) and butterflies & to set seed, had flowering periods extending far beyond
moths (A. foeniculum, Erysimum linifolium). Interestingly, our c. 3-month observation periods. Indeed, E. linifolium
three of the four species particularly attractive to other flowers for approximately 9 months per year in Sussex.
bees are also native to Britain, suggesting that native The attractiveness of such varieties is, therefore, underesti-
plants may be more important for non-Apis and mated in our data.
non-Bombus bees. Closer examination of lavenders showed that hybrid
The factors potentially responsible for variation in L. 9 intermedia varieties were more attractive than both
attractiveness among plant varieties are diverse (e.g. size, L. angustifolia varieties and L. stoechas (Fig. 5). This
shape, colour or scent, reviewed by Pellmyr (2002)). How- difference was not explained by either bloom duration or
ever, as the insects counted were flower-visiting foragers, corolla tube length. In addition, flower colour, which ran-
this variation is presumably largely a result of foraging ged from light (e.g. white ‘Arctic Snow’, ‘Edelweiss’, rose
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
Garden plants for flower-visiting insects 9
Origanum Agastache
1·2 (a) 1·6 (b) Origanum 0·25 (c)
Number of butterflies & moths
1·4
1
Number of other bees
0·2
1·2
per snapshot per m2
Agastache
0·8 Erysimum
1 0·15
Stachys Echium
0·6 0·8
Echium 0·1
0·6
0·4
Achillea 0·4 0·05
0·2
0·2
0 0 0
0 1 2 3 4 0 1 2 3 4 0 1 2 3 4
Number of honeybees (●) or bumble- Number of honeybees (●) or bumble- Number of honeybees (●) or bumble-
bees (○) per snapshot per m2 bees (○) per snapshot per m2 bees (○) per snapshot per m2
Fig. 7. Correlations between the attractiveness of plant varieties to honeybees (●) & bumblebees (○) vs. other bees (a), hover flies (b), and
butterflies & moths (c). The only significant correlation found was between bumblebees and butterflies & moths (r = 0665, P < 0001).
© 2013 The Authors. Functional Ecology © 2013 British Ecological Society, Functional Ecology
10 M. Garbuzov & F. L. W. Ratnieks
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