Entomologia Experimentalis et Applicata 96: 289–298, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

289

Butterfly nectaring flowers: butterfly morphology and flower form

Sarah A. Corbet
Department of Zoology, Downing Street, Cambridge CB2 3EJ, UK
Present address: I St Loy, St Buryan, Penzance TR19 6DH, UK

Accepted: May 23, 2000

Key words: tongue, proboscis, mass, wing loading, nectar, corolla depth, inflorescence, foraging, selectivity

Abstract
The profitability of butterfly foraging depends in part on the corolla depth and clustering of flowers, and the tongue
length, body mass and wing loading of butterflies. Interactions among these attributes of flowers and butterflies
were investigated, using data from a field study in Cornwall and from Porter et al. (1992). The maximum corolla
depth from which a butterfly can feed depends on tongue length, which correlates with the more easily measured
attributes of body mass and wing loading. Small, short-tongued butterflies did not visit deep flowers. The quantity
of nectar sugar per flower necessary for profitable foraging depends on foraging costs, which are expected to
correlate with wing loading. Butterfly species with a high wing loading generally confined their visits to flowers that
were clustered or very nectar-rich. Butterfly species with a low wing loading included solitary and less nectar-rich
flowers in their diet. Body mass and wing loading affect a butterfly’s load-carrying capacity (limiting the distance
between fuelling stops) and cooling rate (limiting the distance between stops for basking or endothermic warming),
and will therefore influence the capacity for floral selectivity and for migration and dispersal. Body mass, wing
loading and tongue length characterised families or subfamilies of butterflies. For example vanessine nymphalids,
with their long tongues and high wing loading, visited the deep, massed flowers of Buddleja davidii, but lycaenids,
with their short tongues and low wing loading, did not. These often visited members of the Asteraceae. Eupatorium
cannabinum, with massed flowers offering abundant and accessible nectar, was visited by butterflies of all tongue
lengths and both high and low wing loading. These findings may help to inform habitat management for butterfly
nectaring flowers.

Introduction on the energetic cost of flight and other foraging activ-

Butterfly species differ in the species of flowers that
they visit (Feber & Smith, 1995). The causal basis
of these differences is poorly understood. Insofaras
nectar consumption increases fitness (Porter, 1992), a
butterfly should select nectaring flowers in a way that
maximises either its energetic efficiency or its net rate
of energy gain, that is, the difference between the gross
rate of gain and the rate of energy expenditure in forag-
ing. The potential gross gain rate depends on the sugar
content of the nectar in each flower (the standing crop)
and the number of flowers emptied per unit time. The
rate of energy consumption during foraging depends
on the forager’s body mass and the mass-specific en-
ergy consumption rate during foraging, which depends
ities and the proportion of foraging time allocated to
each activity. In general, an insect makes a foraging
profit if:
(energy content of mean standing crop per flower) ×
(number of flowers probed per unit time) > (energy
consumption per unit time during foraging).
I therefore consider two attributes of flowers that
govern foraging costs and benefits: corolla depth, and
the extent to which flowers are massed in inflores-
cences.
The standing crop of sugar per flower at a given
time depends on the balance between the secretion
rate, which correlates with flower size (Harder &
Cruzan, 1990), and the rate of depletion by insects.
A deep corolla restricts the nectar clientele to long-
290
tongued species, reducing the rate of depletion and
increasing the probability that a long-tongued visitor
will find a large standing crop in the flower (Cor-
bet et al., 1995; Opler & Krizek, 1984). Among
flower species, the standing crop of nectar generally
varies positively with corolla depth (Harder, 1985;
May, 1988; Prŷs-Jones, 1982). The number of flowers
probed per unit time depends on handling time (de-
termined in part by the relationship between tongue
length and corolla depth; Harder, 1988) and the time
spent travelling between flowers. Interflower travel
times (and costs) are relatively low if flowers are
massed in inflorescences, allowing foragers to move
between flowers by walking, rather than flying. Costs
of travel between inflorescences are low if inflores-
cences are densely crowded.
I consider three attributes of butterflies that gov-
ern foraging costs and benefits: tongue length, body
mass and wing loading (mass per unit area of wing).
The rate of energy consumption during flight is not
well documented for different butterfly species, but it
increases with wing loading among species of moths
(Heinrich, 1993) and euglossine bees (Casey et al.,
1985). If butterfly species with high wing loading
similarly consume energy rapidly, they can forage
profitably only if they have access to large aliquots of
nectar per interflower flight. They should visit flowers
that are massed, allowing them to move from flower to
flower without flying; or flowers that are individually
nectar rich.
In butterflies, tongue length limits the range of
flowers from which nectar can be extracted (Erhardt,
1995; Opler & Krizek, 1984). Similarly, among bum-
blebee species, correlates of flower choice include
tongue length as well as flight costs (Comba et al.,
1999; Corbet, 1997; Corbet et al., 1995; Harder, 1982,
1983; Prŷs-Jones, 1982). Tongue length increases with
body size in some species of flower-visiting insects
(with wing length among species of hawkmoth (Bul-
lock & Pescador, 1983) and with mass within and
among species of bumblebee (Prŷs-Jones, 1982)).
The following predictions are tested in this pa-
per.
(1) In an assemblage of butterfly species, tongue
length correlates with body mass and wing load-
ing.
(2) Butterflies do not forage on flowers with a corolla
depth (or, more precisely, an effective depth to
nectar) that exceeds their tongue length, so that
small butterflies confine their visits to relatively
shallow flowers. Corbet et al. (1995) referred to
this constraint as the depth threshold.
(3) Butterfly species with relatively high flight costs
forage on plant species that give a relatively high
energetic reward per foraging flight. Butterflies
with high wing loading forage mainly on massed
flowers and forage on solitary flowers only if these
are unusually nectar-rich.
Materials and methods
The main study took place in West Cornwall (grid ref.
SW 422233) during August and September 1998. A
few additional records of butterfly feeding were made
at other sites within 20 km: Crean Mill, Lamorna and
Ludgvan. The main study site was small (250 m di-
ameter) and it is assumed that all plant species were
available to all butterfly species. The few records from
other sites involved plant and butterfly species that
were also present at the main site, except for the
Heliotropium hybrid.
For each plant species, I measured the corolla of
ten flowers when possible, taking each from a differ-
ent plant or ramet. Corolla depth was measured from
the most proximal point to which a butterfly might
push the base of its tongue (in sympetalous flowers,
often the base of the deepest cleft in the corolla) to
the supposed location of the nectar at the corolla base.
In Asteraceae, insects probably take nectar from just
above the stamen insertions, where traces of nectar
were sometimes found. I therefore measured from
the base of the deepest cleft in the corolla to the re-
gion where the stamens inserted and the corolla flared
out, at the distal end of the very narrow part of the
corolla tube that is almost filled by the style. In the
tribe Cardueae in particular, the distance measured
was much less than the total corolla length. All plant
species visited by butterflies in this study are peren-
nials (including the monocarpic perennials known as
biennials) except radish (Raphanus sativus), which is
an annual (Table 3). Plant nomenclature follows Stace
(1997).
I found butterflies of 15 species (Table 1). They
represent the following subfamilies: family Pieri-
dae, subfamilies Pierinae (Pieris spp.) and Coliadinae
(Colias); family Nymphalidae, subfamily Vanessinae
(Cynthia, Aglais, Inachis, Polygonia, Vanessa); family
Satyridae (Maniola, Pyronia, Pararge, Lasiommata);
family Lycaenidae, subfamily Lycaeninae (Lycaena,
Polyommatus); family Hesperiidae, subfamily Hes-
 291
Table 1. Tongue length, mass and wing loading index for fifteen species of butterfly found in this study. The table gives mean ± s.e.m.
where appropriate, with the sample size in brackets. Column four shows tongue lengths from Porter et al. (1992)

Species Family Tongue length, Tongue length, Body mass, Wing loading
mm (this study) mm mg index,
(Porter et al., 1992) N m−2

1. Common blue Lycaenidae 8.19 ± 0.20 (5) 8 26 ± 2 (5) 0.66 ± 0.015 (5)
Polyommatus icarus
(Rottemburg)
2. Small copper Lycaenidae 7.34 ± 0.07 (5) 7 34 ± 3 (5) 0.86 ± 0.04 (5)
Lycaena phlaeas (L.)
3. Meadow brown Satyridae 10.45 ± 0.19 (10) 10 70 ± 9 (10) 0.67 ± 0.054 (10)
Maniola jurtina (L.)
4. Gatekeeper Satyridae 7.81 ± 0.12 (9) 8 40 ± 7 (9) 0.58 ± 0.066 (9)
Pyronia tithonus (L.)
5. Speckled wood Satyridae 7.90 ± 0.14 (8) 8 41 ± 4 (8) 0.47 ± 0.028 (8)
Pararge aegeria (L.)
6. Small white Pieridae, 12.18 ± 0.14 (5) 15 67 ± 6 (5) 0.56 ± 0.040 (5)
Pieris rapae (L.) Pierinae
7. Large white Pieridae, 14.72 ± 0.39 (10) 16 128 ± 9 (10) 0.83 ± 0.039 (10)
Pieris brassicae (L.) Pierinae
8. Green-veined white Pieridae, 10.53 ± 0.47 (4) 11 56 ± 5 (4) 0.59 ± 0.052 (4)
Pieris napi (L.) Pierinae
9. Clouded yellow Pieridae, 14.09 ± 0.21 (5) 15 96 ± 5 (5) 0.79 ± 0.026 (5)
Colias croceus Coliadinae
(Geoffroy)
10. Small skipper Hesperiidae 15.00 (2) 14 61 (2) 2.34 (2)
Thymelicus sylvestris (Poda)
11. Comma Nymphalidae 12.94 ± 0.98 (2) 12 192 (2) 1.63 (2)
Polygonia c-album (L.)
12. Painted lady Nymphalidae 15.10 ± 0.31 (4) 14 218 ± 29 (4) 1.54 ± 0.224 (4)
Cynthia cardui (L.)
13. Red admiral Nymphalidae 15.11 ± 0.28 (9) 13 276 ± 28 (9) 1.71 ± 0.151 (9)
Vanessa atalanta (L.)
14. Small tortoiseshell Nymphalidae 14.17 ± 0.42 (5) 14 136 ± 13 (5) 1.20 ± 0.095 (5)
Aglais urticae (L.)
15. Peacock Nymphalidae 16.19 ± 0.41 (3) 17 275 ± 31 (3) 1.60 ± 0.112 (3)
Inachis io (L.)

periinae (Thymelicus). Butterfly nomenclature follows
Dennis (1977), Thomas & Lewington (1991) and
Pullin (1995).
Many studies of foraging insects involve regular
systematic counts of visits and of flowers in a limited
area (e.g., Corbet et al., 1995). Counting visits and
flowers was not practicable here because butterflies
were so sparse that it was necessary to monitor a very
large area. Selectivity is revealed by differences be-
tween butterfly species in flowers visited, rather than
by comparing counts of flowers visited and flowers
available.
Flower visits were recorded only if a butterfly def-
initely probed a flower rather than just resting on it,
and they usually involved sustained feeding. Even
so, some recorded interactions may represent one-off
sampling visits rather than regular associations, be-
cause I did not quantify visit frequency. My findings
are compared with those of Porter et al. (1992), who
list flower-visiting records of British butterfly species.
They do not state that their list is limited to vis-
292

its involving sustained probing rather than sampling,

settling or basking.
To measure butterflies, I captured them and within
a few hours (usually two) weighed them to the near-
est 2 mg on an Ohaus CT portable electronic balance.
When possible, I sampled five individuals of each sex,
but for some species fewer individuals were available
(Table 1). The head was removed and the proboscis
was unrolled and measured with Mitutoyo digital
calipers (Mitutoyo (UK) Ltd, Andover, UK) from the
tip to the front of the compound eye. Occasionally a
proboscis was distorted and apparently nonfunctional,
and could not be measured.
Wing loading in N m−2 is mg/S where m is
the body mass in kg, g is the gravitation constant
(9.80665 m s−2 ), and S is the effective area of the
wings during flight, in m2 . Wing areas of all four
wings were measured using a Delta T leaf area meter
(Delta T, Cambridge, UK) and summed for each spec-
imen. To the extent that wings overlap during flight,
this summed area measurement overestimates the ef-
fective wing area, resulting in underestimation of wing
loading. I therefore refer to this underestimate of wing
loading as the wing loading index. Any damaged wing
was assumed to have the same surface area as its pair.
If both wings of a pair were incomplete, I did not use
the wing area measurement.
Statistical analysis was performed using SPSS 9.0.
The data set consisted of measurements for up to
ten individuals of each of 15 species. Analysis was
based, not on data for individual butterflies, but on Figure 1. Mean measurements for fifteen species of butterfly be-
species means or subfamily means. A Kolmogorov- longing to five families: Satyridae (white circles); Pieridae (black
triangles); Nymphalidae, Vanessinae (white triangles); Lycaenidae
Smirnov test showed that all variables were normally
(black diamonds); and Hesperiidae (black dot), showing (a) tongue
distributed after log transformation of body mass and length versus log body mass, and (b) wing area versus log body
wing loading index. Preliminary analysis of variance mass.
showed that any effect of sex on the measured at-
tributes of the butterflies, whether log transformed or
tongue length correlated significantly (P<0.001) with
not, was not significant at the 5% probability level, so
log body mass, wing area and log wing loading index
the sexes were pooled for further analysis.
(Table 2). Omission of the mean for the small skip-
per improved the correlation of tongue length with log
body mass and wing area, but not with log wing load-
Results
ing index (Table 2). The significant correlations were
not all due to inclusion of several species in each sub-
Mean values of tongue length (Figure 1a) and wing
family. When I used the means of all the species means
area (Figure 1b) of the 15 species increased with body
in each subfamily, again omitting the skippers, tongue
mass. The small skipper (Thymelicus sylvestris), the
length correlated significantly with log mass and wing
sole representative of the Hesperiidae, had a longer
area but not with log wing loading index (Table 2).
tongue and a smaller wing area relative to its body
No butterflies were seen to probe flowers deeper
mass than the other species (Figures 1a, b). With
than the mean tongue length of their species (Ta-
each species mean treated as an independent data
ble 3). Potential interactions between a butterfly and a
point (a ‘naïve’ interspecific analysis; Grafen, 1989),

Table 2. Pearson correlation coefficients for tongue length versus log body mass,
total wing area and log wing loading index, based on species means of 2–10
individuals in each of 15 species, and means of the species means in six subfam-
ilies (total: 86 individuals). Correlations are improved by omitting the skippers
(Hesperiidae, Hesperiinae), leaving 14 species in 5 subfamilies
Log mass
All species (n = 15) 0.862∗∗∗
Species, skipper 0.933∗∗∗
omitted (n = 14)
Subfamilies, skippers 0.932∗
omitted (n = 5)
∗ P < 0.05; ∗∗ P < 0.01; ∗∗∗ P < 0.001
Figure 2. Relation between the mean effective corolla depth of the
plants visited by 15 butterfly species and the butterflies’ mean wing
loading index (N m−2 ), for plant species with solitary (black dots)
or massed (open circles) flowers. Each point represents at least one
visit. The line marks a wing loading index of 0.9 N m−2 . Butterflies
with wing loading greater than this generally confined their visits to
massed or nectar-rich flowers.
flower with a corolla longer than its tongue are marked
X in Table 3. I saw no such interactions. Although
plants with deep corollas were visited by only the
few long-tongued butterfly species, some effectively
short-tubed plants such as Centaurea nigra and Eupa-
torium cannabinum received visits from a wide range
of butterfly species.
Pierids, satyrids and lycaenids all had wing load-
ing indices below 0.9 N m−2 , and visited both massed
and solitary flowers. The vanessine nymphalids and
the hesperiid all had wing loading indices exceeding
1.2 N m−2 , and did not generally visit solitary flow-
ers (Figure 2). The exception was a visit by a red
admiral (Vanessa atalanta) to bramble (Rubus fruti-
293
Wing area Long wing loading index
0.672∗∗ 0.733∗∗
0.933∗∗∗ 0.724∗∗∗
0.935∗ n.s.
cosus agg.), which produces large and often clustered
flowers that can be nectar-rich. The standing crop of
nectar sugar in flowers of a given species fluctuates
markedly over time (Corbet et al., 1995), but that of
R. fruticosus is often so large that this has been de-
scribed as a cornucopia species sensu Mosquin (1971)
(Yeboah Gyan & Woodell, 1987a,b). The only single-
flowered species with a long corolla was red campion
(Silene dioica). Seven butterfly species have tongues
longer than the corolla of this species, five with wing
loading indices greater than 1.2 N m−2 (four vanes-
sine nymphalids and a hesperiid) and two pierids with
wing loading indices below 0.85 N m−2 . Only the two
species with low wing loading (Coleus croceus and
Pieris brassicae) probed red campion flowers.
Discussion
In this assemblage of butterfly species, tongue length
correlates with body mass and wing loading, support-
ing hypothesis 1.
My observations support hypothesis 2, in that but-
terflies did not feed on flowers with a mean corolla
depth greater than their mean tongue length. Hypoth-
esis 2 is also supported by the findings of Porter et al.
(1992), with two apparent exceptions. These authors
recorded two butterfly species, small whites (Pieris
rapae) and green-veined whites (P. napi), visiting red
campion (Silene dioica), for which my mean corolla
length measurement (13.30 mm in female flowers) ex-
ceeds the mean tongue length I measured for small
whites (12.18 mm) and green-veined whites P. napi
(10.53 mm). Given the variability in the lengths of
both tongues (small whites, range 12.01 to 12.72 mm
(n = 5); green-veined whites, range 9.30–11.51 mm,
n = 4) and flowers (range 11–15 mm, n = 10), some
Table 3. Plant species on which at least one butterfly visit was recorded in this study, with corolla depth (mean ± s.e.m.) and flower attributes. Column 5 shows which flowers
294

are massed in inflorescences, and column 6 shows which are members of the important family Asteraceae. Flowers present also in the list of Porter et al. (1992) are in bold.
Butterfly visits in that list are shown as asterisks. Plant and butterfly species in the list of Porter et al. but not in my list are omitted. Potential butterfly/flower combinations
in which the corolla depth would exceed the tongue length are marked X

mean corolla depth, mm

± s.e.m.
no. flowers measured
no. plants [or ramets]
are flowers massed?
Asteraceae
no. butterfly spp. (this study)
no. butterfly spp. (Porter)
Lycaena phlaeas
Pyronia tithonus
Pararge aegeria
Polyommatus icarus
Maniola jurtina
Pieris napi
Pieris rapae
Polygonia c-album
Coleus croceus
Aglais urticae
Pieris brassicae
Thymelicus sylvestris
Cynthia cardui
Vanessa atalanta
Inachis io

Mean tongue length, mm 7.34 7.81 7.90 8.19 10.45 10.53 12.18 12.94 14.09 14.17 14.72 15.00 15.10 15.11 16.19

Rubus fruticosus agg. 0.00 0 6 9 Y∗ Y∗ Y∗ Y ∗ ∗ Y ∗ Y

Jasione montana 0.00 0 Y 1 Y
Escallonia sp. nr. macrantha 0.00 0 1 Y
Geranium sp. nr. endressii 0.00 0 1 Y
Taraxacum officinale agg. 0.33 0.03 10 2 Y Y 1 7 ∗ ∗ ∗ Y∗ ∗ ∗ ∗

Sedum spectabile 2.05 0.06 10 2 Y 6 Y Y Y Y Y

Hebe cv. A 2.69 0.07 10 1 Y 1 Y
Eupatorium cannabinum 2.71 0.08 10 10 Y Y 8 10 Y Y Y Y Y ∗ Y Y Y∗ ∗

Leucanthemum × superbum 2.99 0.10 10 [10] Y Y 5 Y Y Y Y

Erigeron karvinskianus 3.31 0.04 10 [10] Y Y 6 Y Y Y Y Y Y
Senecio jacobaea 3.72 0.09 10 10 Y Y 6 9 Y Y Y Y ∗ ∗ Y ∗

Hebe cv. B 3.93 0.08 10 1 Y 2 Y Y

Mentha aquatica 4.11 0.16 10 10 Y 6 8 ∗ Y Y Y Y∗ Y Y ∗

Heliotropium hybrid 4.20 0.10 10 1 Y 4 Y Y Y

Origanum vulgare 4.25 0.17 10 10 Y 2 8 Y ∗ ∗ Y ∗ ∗ ∗ ∗

Centaurea nigra 4.79 0.23 10 10 Y Y 11 12 Y∗ Y∗ Y Y ∗ Y∗ ∗ Y∗ Y∗ Y∗ Y Y

Ligustrum sp. 5.65 0.19 2 2 Y 1 9 ∗ ∗ ∗ ∗ ∗ Y ∗ ∗ ∗

Buddleja globosa 6.01 0.16 10 1 Y 3 Y Y Y

Geranium robertianum 6.12 0.19 10 10 2 Y Y
Prunella vulgaris 7.22 0.22 10 10 Y 3 4 ∗ Y Y
Agastache urticifolia 7.57 0.13 10 1 Y 3 X Y Y Y
Raphanus sativus 7.69 0.13 10 10 1 X Y
Buddleja davidii 9.20 0.22 10 10 Y 5 X X X X Y Y Y Y Y
Agapanthus praecox 12.53 0.23 10 10 Y 3 X X X X X X X Y Y Y
Silene dioica 13.30 0.42 10 10 2 4 X X X X X X∗ X∗ X Y Y

individuals of both pierid species could reach the nec-
tar in at least some S. dioica flowers. Thus hypothesis
2 is supported without exceptions.
Hypothesis 3, that butterfly species with high wing
loading forage mainly on flowers that are massed or
unusually nectar-rich, is supported by my findings,
and by the subset of those of Porter et al. (1992) in-
volving the same species, in that no high-wing-loading
species fed on solitary flowers other than the nectar-
rich bramble (Rubus fruticosus). However, the total
list of Porter et al. includes apparent exceptions among
species not represented in my study. Eight of the 38
butterfly-visited plant species that they examined have
solitary flowers. As in my study, Rubus fruticosus
received visits from many butterfly species including
six nymphalids and two hesperiids, and red campion
(Silene dioica) was visited only by pierids. The re-
lated ragged robin (Lychnis flos-cuculi), with a shorter
corolla and a higher midday nectar secretion rate than
red campion (Comba et al., 1999), was visited not
only by six pierid species but also by three species
of hesperiids, one satyrid and two species of fritillary.
Fritillaries are nymphalids of the subfamilies Argyn-
ninae and Melitaeinae, for which the wing loading
index has not been measured but is probably lower
than that of the vanessine nymphalids considered here.
Fritillaries were the only nymphalids that visited tor-
mentil (Potentilla erecta) and buttercups (Ranunculus
species), but cuckoo flower (Cardamine pratensis) and
greater stitchwort (Stellaria holostea) were visited by
vanessine nymphalids (a peacock Inachis io and a
small tortoiseshell Aglais urticae respectively). In the
list of Porter et al. (1992) the only apparent exceptions
to hypothesis 3 are these and buttercups visited by
hesperiids. Those visits may have involved sampling
rather than sustained probing. Thus my findings sup-
port hypothesis 3 and those of Porter et al. probably
do not falsify it.
The assemblage of attributes that influence flower
choice also influence other aspects of butterfly biol-
ogy. Attributes that influence flower choice by af-
fecting flight costs and rate of energy intake will, in
conjunction with fuel capacity, determine how far a
butterfly can fly between stops. This has implications
for flower selectivity and migration, both of which de-
pend on both the potential duration of uninterrupted
flight (determining how far an insect can fly without
stopping to warm up), and load-carrying capacity (de-
termining how far an insect can fly between refuelling
stops).
295
The potential duration of uninterrupted flight de-
pends on maintenance of a high enough body tem-
perature for flight. Body mass is positively associated
with the capacity for endothermy in bees (Stone &
Willmer, 1989) and probably in butterflies as well
(Heinrich, 1993). Small insects have relatively high
rates of convective cooling and tend to cool during
flight. Heinrich (1993) has suggested that at low am-
bient temperatures, insects weighing less than about
200 mg probably cannot warm up metabolically fast
enough to achieve flight temperature or to maintain or
elevate thoracic temperature during flight. Small in-
sects can extend flight duration by pre-flight basking to
elevate the thoracic temperature well above the flight
threshold, but they must stop flying when the thoracic
temperature falls below this threshold. Hence at low
ambient temperatures, only butterflies weighing more
than about 200 mg can maintain continuous uninter-
rupted flight (Heinrich, 1993). Presumably smaller
species can migrate when the ambient temperature
(or, more precisely, a black globe temperature that
allows for radiative warming in the sun: Pivnick &
McNeil, 1987) equals or exceeds the flight threshold.
Of the species considered here, nine (all the pierids
and vanessine nymphalids) are classified as migrants
by Thomas (1984 and pers. comm.). Only three of
these weigh more than 200 mg. Presumably the lim-
ited endothermic capacity of the other six allows them
to sustain uninterrupted flight only during warm sunny
weather.
Sustained flight also depends on the amount of fuel
carried. Load-carrying capacity depends on the ratio
of flight muscle mass to total body mass. This ratio
is expected to be high in species that frequently need
maximal power output for lifting loads such as large
nectar meals or batches of unlaid eggs (Marden, 1987).
The ability to sustain flight has implications for
the population dynamics and conservation prospects
of butterflies. The relation between mobility and mor-
phology is illustrated in Figure 3. With the exception
of the hesperiid, all the species considered here that
Thomas (1984 and pers. comm.) classifies as seden-
tary species with closed populations have tongues less
than 10.5 mm long and a wing loading index below
0.9 N m−2 . They appear in the lower left quadrant
of Figure 3. All the species that Thomas classifies
as migratory or having open populations have longer
tongues, and the vanessine nymphalids (but not the
pierids) also have a higher wing loading index.
The ability to sustain flight also has implications
for flower selectivity. An insect capable of making
296

Figure 3. Mean tongue length of 15 butterfly species, with family symbols as in Figure 1, plotted against wing loading index. Line A at
0.9 N m−2 and line B at 10.5 mm delimit a lower left quadrant in which all species (1–5 in Table 1) form closed populations (Thomas, 1984 and
pers. comm.). Above line B, all species (6–9 and 11–15) except the small skipper (10 in Table 1) are classified as forming open or migratory
populations (Thomas, 1984 and pers. comm.).

long flights and carrying heavy loads can be highly
selective, visiting only nectar-rich flowers or inflo-
rescences, and taking meals that are large and few.
An insect capable only of brief flights and of car-
rying light loads might be less selective, bypassing
fewer flowers when foraging, and taking smaller, more
frequent meals. May (1992) found a difference in se-
lectivity between two syntopic butterfly species with
mean body weights of 195 and 231 mg respectively
(May, 1988). The heavier species, Phoebis sennae L.
(Pieridae), achieved a higher net rate of energy gain
than the lighter Agraulis vanillae L. (a non-vanessine
nymphalid) by selecting flowers with more nectar.
May (1988) asked why the smaller species did not
forage as profitably as the larger species. Perhaps the
paradox of its lower selectivity and hence apparently
suboptimal foraging can be interpreted in terms of
thermoregulatory and load-carrying constraints.
Nearly all the plant species that butterflies visited
for nectar flowers are perennials. Presumably, few an-
nuals contain enough nectar to make a visit profitable.
Feber & Smith (1995) and Dover (1996) found that
butterflies visited perennials more frequently than an-
nuals in relation to their relative abundance, as do
bumblebees (Fussell & Corbet, 1992a, b). In the total
list of Porter et al. (1992) all 38 plant species visited
by butterflies were herbaceous or woody perennials,
not annuals. In my study the only annual visited was
radish (Raphanus sativus, Brassicaceae). The related
Sinapis arvensis is visited by Pieris napi, P. rapae and
P. brassicae on conservation headlands in arable fields
(Dover, 1989). On annually-ploughed land, which in-
evitably yields few perennials, Sinapis is one of a
small group of annuals rich enough in nectar to permit
butterflies to forage profitably, and even this is appar-
ently useful only for pierids with their combination of
a long tongue with a moderate wing loading.
For an assemblage of butterfly species in Britain
in late summer, this paper has shown a relationship
between the tongue length and wing loading index of
a butterfly species and features of the flower species
that it probes for nectar. Although many of the plant
species studied here are exotic, the generalisations de-
rived from them are broadly supported by comparison
with the more comprehensive set of records of Porter
et al. (1992), and may be expected to illuminate flower

choice by butterflies in seminatural habitats as well as
gardens.
These generalisations may help inform the man-
agement of vegetation for butterflies. In general,
species with shorter tongues, notably lycaenids and
some satyrids, do not visit deep flowers (including the
well-known butterfly flower Buddleja davidii). Provi-
sion for these shorter-tongued species should include
Asteraceae such as Eupatorium, Senecio and Leucan-
themum. Deep flowers, with their potentially abundant
nectar, were much visited by butterflies with tongues
long enough to reach to the corolla base. Pierids,
which combine a relatively long tongue with a mod-
erately low wing loading (Figure 1b), can include in
their diet deep solitary flowers such as Silene dioica
and even annual brassicaceous arable weeds. The
vanessine nymphalids, long-tongued butterflies with a
high wing loading, usually visited flowers massed in
dense inflorescences, such as Buddleja davidii, Hebe
species and Asteraceae of the tribe Cardueae. Proba-
bly they cannot forage profitably on solitary flowers
unless, like Rubus fruticosus, these are very rich in
nectar. Many important nectaring plants are members
of the family Asteraceae, and Eupatorium and nectar-
rich Cardueae such as Centaurea nigra host large
numbers of butterfly species spanning a wide range of
morphological types.
Acknowledgements
I thank Andreas Erhardt and Jeremy Thomas for pro-
viding unpublished data and ideas, Andrew Barron,
Charlie Ellington, Lawrence Harder and Robin Woot-
ton for comments on an earlier draft, Paul Freeman
for help with equipment and John Hobart and Philip
Corbet for help with field recording.
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